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CICHLIDS yearbook

Ad Konings (Ed.)

Volume 1The

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Cover photographs:1 - Benthochromis tricoti, Lake Tanganyika, Zambia.2 - “Cichlasoma” labridens, Media Luna, Mexico.3 - Symphysodon aequifasciatus, variety.4 - Otopharynx lithobates, Zimbawe Rock, Lake Malawi, Malawi.

Text and photographs by Ad Koningsexcept as otherwise indicated

Mary Bailey (Crediton, UK)corrected the manuscript

The editor wants to thank the following persons whosupplied various cichlids for photographic purposes:Peter Baasch (Stegen, Germany)Marc Danhieux (Maltavi, Hohenahr-Erda, Germany)René Krüter (Krüter Tropicals, Rotterdam, Netherlands)Roland Numrich (Mimbon Aquarium, Köln, Germany)Edwin Reitz (Aquapport, Ronnenberg, Germany)Dirk Verduijn (Verduyn Cichlids, Zevenhuizen, Netherlands)

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Distributors:

USA: Old World Exotic Fish, Inc., P.O.Box 970583, Miami, Florida 33197UK: Finz (U.K.), Ltd., Lady Ann Mills, Lady Ann Road, Batley, West Yorkshire WF17 0PSSweden: Fohrman Aquaristik AB, Pepparplan 4, 393 65 KalmarGermany: Aquapport (Edwin Reitz), Köselstraße 20, 3003 RonnenbergNetherlands: NVC, Lieshoutseweg 31, 5708 CW Stiphout

ISBN 3-928457-00-4

Copyright © 1991. Verlag Dr. Gertrud Dudin. All rights reserved.No part of this publication may be reproduced, stored in a retrieval system, or transmitted in any form or by any means—electronic, mechanical, photocopying, recording or otherwise—without the prior permission of the authors, and the pub-lisher.

Edition Cichlid PressVerlag Dr. Gertrud Dudin, 6837 St. Leon-Rot, Germany

Printed by RAKET B.V., Pijnacker, Holland

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Fascination with cichlids has taken many aquariststo the point where they go and observe these won-derful fishes in the natural habitat. The hunger formore knowledge about the cichlids’ behavior anddistribution, and especially for more new specieshas led to the introduction of an overwhelming ar-ray of previously unknown species and varieties.The speed with which new species are discoveredor re-discovered is sometimes too great to give anewcomer a chance to establish itself in the hobby.We, the authors of the yearbook, want to give achance to those species which are new or unknownand which deserve a wide distribution amonghobbyists. Starting with this yearbook every yearthe newest, the most important, the most interest-ing and, not least, the most beautiful cichlids willbe introduced to the hobbyist. All (re-)introductionswill be accompanied by a color picture and the cur-rently known information. Such information canonly be supplied by aquarists (or scientists) whoare specialists in their fields. Many authors havecontributed to the first edition of the cichlid year-book. They will be introduced by the editor.

The yearbook starts with the section “Tangan-yikan Cichlids”. René Krüter, who is importer ofTanganyikan cichlids (Krüter Tropicals, Holland),has made several expeditions to Lake Tanganyika.He reports on his observations of Benthochromistricoti and some other species.

The second section, “Malawian Cichlids” is high-lighted by an article written by Dr. EthelwynnTrewavas. She is the world’s most renowned cichlid-ichthyologist and has worked for more than 60 yearson Malawi cichlids.

Photographer, diver and naturalist Mark Smith(Fountain Valley, California) reflects his vivid in-terest in cichlids in the various articles he has con-tributed to the yearbook. Peter Baasch (Freiburg,Germany) has bred several Malawi cichlids for thefirst time in captivity. He reports on a beautifulpredator from the lake.

Victoria cichlids deserve a much better apprecia-tion among aquarists than has been the case up tonow. Laif DeMason, who imports the widest vari-ety of cichlids from all over the world (Old WorldExotic Fish, Miami, Florida), has made several ex-peditions to Lake Victoria and knows these fish from

The CICHLIDS yearbook

first hand. His contribution deals with beautiful andnew discoveries.

The cichlids of West Africa are totally differentfrom the previous groups. Specialist RolandNumrich (Mimbon Aquarium, Köln, Germany) hasmade many expeditions to this part of the world andhas caught, bred and raised many of the newer finds.His article describes two new species of Chromido-tilapia.

The section “Central American Cichlids” startswith an excellent article by Juan Miguel ArtigasAzas (San Luis Potosí, Mexico), who has more than15 years experience of the natural habitats of Mexi-can cichlids. On one of his countless expeditionshe re-discovered Paraneetroplus nebuliferum,which had been elusive since its original descrip-tion in 1860.

Willem Heijns (Stiphout, Netherlands) is editorof the periodical of the Dutch Cichlid Associationand has kept and bred almost every Central Ameri-can cichlid. He has contributed several fine reportson the latest developments.

The section “South American Cichlids” has beenwritten by two esteemed specialists, Ron Bernard(Rotterdam, Netherlands) and Frank Warzel (Mainz,Germany). Ron has written many articles dealingwith South American cichlids and has collectedcichlids in Peru. Frank is super-specialized onCrenicichla and knows more about the pike cichlidsthan anybody else.

Gerard Tijsseling (Gouda, Netherlands) and JohnSzwechlowicz (Holton-Le-Clay, UK) provide uswith valuable tips on how to better enjoy and breedcichlids.

The yearbook ends with a literature section wherea few very important publications, related tocichlids, are discussed. Nobody else could do thisbetter than the two authors Martin Geerts (Swalmen,Netherlands) and Lee Finley (Pascoag, Rhode Is-land). Martin is the ichthyological conscience of theDutch Cichlid Association and Lee that of theAmerican Cichlid Association.

This fantastic team of authors provides for an au-thoritative treatment of the latest developments inthe cichlid-keeping hobby.

Your editor

Ad Konings (Editor)

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In 1948 Poll described Benthochromis tricoti asHaplotaxodon tricoti, but later, in 1984, he split thetwo different populations, which he originallythought belonged to one species, into two species:Haplotaxodon tricoti and H. melanoides. Two yearslater he gave these two species their own genus,Benthochromis.

The name Benthochromis alludes to the assump-tion that the species of this genus live at deep lev-els. Benthos is Greek for “depth”. B. tricoti differsfrom B. melanoides by its larger maximum size

TANGANYIKANCICHLIDS

The giant featherfin

René Krüter

A courting male Benthochromis tricoti, taken in Zambian waters by René Krüter.

which is about 20 cm; B. melanoides remains about3 cm smaller. Also B. tricoti, at least in males, haslongitudinal stripes on the body. B. melanoides hasa number of black blotches on the head. The fe-males of both species have an overall silvery-graycoloration.

The food of B. tricoti consists of small inverte-brates, such as crustaceans and zooplankton. Al-though it attains a considerable size B. tricoti is nota piscivore.

B. tricoti is mainly found in the southern part of

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A male Benthochromis tricoti displaying.

When courting, Benthochromis tricoti has its mouth wideopen. This is also seen in species of Cyprichromis.

Lake Tanganyika. It is abundant in the deep to verydeep layers. Several investigations, including thosedone by the Zambian Fisheries, revealed that thedensest populations were to be found between 100and 150 meters deep. Another species which isabundant at these levels is Cyphotilapia frontosa.

Normally I found B. tricoti where the rock for-mations on shore descended at a steep angle. I havenever found this species in water shallower than 35meters and at this level usually no more than a fewsolitary individuals. At 65 meters I found manymore individuals, including several territorial males.The population density between 100 and 150 me-ters may be so high that many escape to higher lev-els or they may migrate to higher levels to spawn.Unfortunately it is impossible, with the availablediving equipment, to descend to deeper levels insafety.

Sexually active males try to obtain a territory ontop of a conspicuous rock and defend this, likeOphthalmotilapia ventralis, against conspecifics. Ihave, however, never found sand in these territo-ries. The bare rock is defended against rivals. Dur-ing territorial disputes males show their finest col-oration; the spectacle of fighting and courting B.tricoti is one of the most fascinating splendors ofthe lake.

The courting behavior is very characteristic. Assoon as a female approaches the breeding colony,she is surrounded by a few males. Each of them

tries to lead her — while swaying his body heavily— to his rock. When the male arrives at the rock hetakes a peculiar posture. With all fins erect the maleBenthochromis puts himself in an oblique, head-upposition. The dorsal fin is fully erected so that thefirst few spines point forward. Also the mouth isopened completely and the buccal sac is inflated.The male remains in this position for a few secondsshowing his best to the female. Sometimes a weakvibration enhances the display.

Unfortunately I have never witnessed a spawn-ing. The size of the fry in the mouth of some fe-males is remarkable. Sometimes the fry had a lengthbetween 3 and 4 cm. This may indicate that the lar-vae and fry feed inside the female’s mouth. If theyfeed within the mouth the female has to eat appro-

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A male Benthochromis tricoti leading a female (not visible) to its nest.

priate food, or she must release the fry from time totime to let them eat by themselves. This may fur-ther indicate that the female takes care of the fryeven after they have been released for the first time.Successful breeding in captivity may give an an-swer to these questions.

B. tricoti is an ideal aquarium resident. It is nei-ther aggressive nor predatory. Despite its relativelylarge dimensions, B. tricoti does not require an enor-mous aquarium. After a relatively short acclimati-zation period males regain their splendid colora-tion. This in contrast to e.g. Cunningtonia andCyathopharynx, who need a long period of accli-matization before full coloration returns.

B. tricoti is one of the most difficult to catch fishin Lake Tanganyika. The main reason is the depthof their preferred habitat. At just a few known loca-tions they can be found at depths of less than 40meters. Yet 40 meters is rather deep for a Scubadiver. If we want to collect fish as well we encoun-ter some problems. A method has to be developedto bring these fish to the surface alive. It is fatal totake a cichlid straight from these depths to the sur-face. The closed swim-bladder, which is filled withair, will expand drastically because of the decreasedpressure at the surface and may even burst. The

other internal organs will be forced out via the anusand throat, which leads to the inevitable death ofthe fish. That is why cichlids from these depths haveto be decompressed very carefully, in order to givethe blood enough time to absorb the excess air fromthe bladder. The time it takes to surface B. tricoti,caught at 35 meters, is five days.

I have noticed that fish can voluntarily alter theirdepth range much quicker than when they are forcedto do it. My observation is that decompression takesmuch longer when a fish is brought to the surfaceagainst its will.

Another problem with B. tricoti is the females.When we descend to a depth of 40 meters allBenthochromis will be scared away. The territorialmales will return after a while, but of the femalesno sign at all. The females seem to stay in smallschools far away from anything dangerous. The fewfemales that have been caught up to now got acci-dentally stuck in the net.

Personally speaking, I think B. tricoti is one ofthe most beautiful aquarium fishes. It has a veryinteresting behavior and I hope to solve a few of itsmysteries on my future trips to the lake.

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The cichlids from Lake Tanganyika are well-knownfor their diverse behavior and enjoy a wide interestamong hobbyists. Of particular interest are the so-called sand-cichlids. These fish, mainly of the ge-nus Xenotilapia, live over the open sandfloor of thelake. To protect them against predators their bodiesare silvery or sand colored and they live in largeschools, which gives protection to the individual.During the breeding season the males of at leastthree sand dwelling species acquire a beautiful andconspicuous breeding dress. One of these species,Enantiopus sp. “Kilesa” has not been described yet;the other two species are Enantiopus melanogenysand Xenotilapia ochrogenys. The latter two specieshave been observed in the natural habitat in Burundi,Tanzanian, and Zambian waters, but are reportedlyalso present along the western coast of the lake(Poll, 1956). E. sp. “Kilesa” was exported fromZaire, probably from a location named Kalumbie.

E. melanogenys was described by Boulenger in1898 as Ectodus melanogenys. A few years later, in1906, the same author placed it in Enantiopus. Pollplaced this cichlid in Xenotilapia (1951) and backin Enantiopus (1986). X. ochrogenys was describedas Enantiopus ochrogenys by Boulenger in 1914 andplaced – by Poll (1946) – in Xenotilapia.

A male Enantiopus sp. “Kilesa” courting a female.

Three beautiful sanddwelling cichlids

Greenwood (1978) demonstrated clearly that boththese species possess different anatomical featureswhich means that they are unlikely to belong to thesame genus.

E. melanogenys and E. sp. “Kilesa” are very slen-der, elongated cichlids with a long snout. E. sp.“Kilesa” differs from E. melanogenys by the shorterlower jaw (see photos). All known species of thegenus Xenotilapia have slender bodies but roundedheads. Their snout profiles descend steeply. The twospecies of Enantiopus reach a maximum size ofabout 16 cm; the maximum size of X. ochrogenysis about 12 cm.

There are no distinct geographical races knownfor E. melanogenys nor for E. sp. “Kilesa”. Thepopulations of X. ochrogenys along the coast ofBurundi and Tanzania do not vary – regarding malecoloration – but in Cameron Bay, Zambia, anothervariety (or species) has been observed. The so-called “Ndole Ochrogenys” differs from the wide-spread variant of X. ochrogenys by prominent blackspots on the flanks and by the larger maximum size– approximately 14 cm.

These sanddwelling cichlids forage in largeschools in rather shallow water. The maximumdepth recorded for E. melanogenys is 40 m but most

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A male Enantiopus melanogenys defending its territory.

The lower jaw is shorter in E. sp. “Kilesa” (above) than in

specimens are seen in depths of less than 10m. Whennot breeding males have the same sandy, silverycoloration as females. It gives them optimum cam-ouflage on the sandy bottom. Most individuals ofone school remain together in the same schoolthroughout their lives. A school is probably formedat the moment of the simultanuous release by themouthbrooding females. The youngsters grow andbreed together until they die after about three years.

All three species are mouthbrooders in whichonly the female incubates the eggs and larvae.Breeding has been observed throughout the year butthe highest activity takes place during the rainy sea-son – from December until May. During the rest ofthe year most of the schools move around and for-age in different areas. X. ochrogenys normally oc-curs in small schools of up to 30 members, but theschools of the two Enantiopus may number in thehundreds. Because there are so many fish in oneschool together they do not have to seek shelterwhen they want to breed, as many other sand-dwelling cichlids do. Since most members of theschool have the same age they all attain their re-productive phase at about the same time. It seemsthat X. ochrogenys has attached itself to the breed-

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The territory of E. melanogenys is flat (above) whereas

A male Xenotilapia ochrogenys from the Ndole Bay population.

ing colonies of the Enantiopus since they are usu-ally found breeding together.

During the breeding season males defend theirterritories and try to persuade females to spawn intheir nests. The actual breeding may take place inbouts which would last for less than a week. Dur-ing this period the fish do not eat but concentrateon spawning. The season begins when the malesstart staking out their territories in the sand. A maleE. melanogenys digs a flat, saucer-shaped territorywith a diameter of about 60 cm. In the center of theterritory, he digs a small pit with a diameter of about15 cm. This will be the nest in which the spawningwill take place. The territory of X. ochrogenys israther peculiar; it consists of three to eight turretsbuilt by heaping sand. They are put in a circle aroundthe spawning-site which is a round, saucer-shapedpit with a diameter of approximately 10 cm. Theterritory of E. sp. “Kilesa” is a very interesting“mixture” of the former two types. In an aquarium,a male will occupy a large area in which he makesseveral nests! These nests are shallow pits with adiameter of about 15 cm. Around these nests themale heaps sand turrets all over its territory, some-times more than 20. A spawning usually takes place

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in a single nest but next time they may spawn inanother.

The nests of these sanddwelling cichlids are me-ticulously cleared of small pebbles which are largerthan the prevailing sandgrains. The reason whymales take the trouble to remove larger grains isobvious: the female will not then mistake a tiny peb-ble for an egg when she collects them during spawn-ing. On an evenly structured sand-nest the eggs aremore conspicuous and can be collected faster.

When the arena is ready the male starts courting.Here we notice some differences between Enan-tiopus and Xenotilapia. A male X. ochrogenys at-tracts a female to the nest with its fins fully ex-tended, except the first part of the dorsal fin whichis just half-extended. A male Enantiopus attractsfemales in a most peculiar fashion. Rivals are chasedwith all fins erect, but females are seduced to thenest with all fins down! The only thing extended isthe buccal cavity as in Xenotilapia. Furthermore themale lies almost completely on its side when at-tracting the other sex. When more than one male isaround the female is not chased when she ignoresthe courtship of a male. As a female passes over thedifferent territories we see the males, one by one,lying on their sides and trying to seduce her. Themales, however, stay in their territories.

When a female is ready to spawn she responds tothe display of the male. As soon as she enters thenest, the male circles around her with fins erect –in Enantiopus as well as Xenotilapia. Males are soexcited that they first chase all other fish away fromthe nest. Meanwhile, the female remains motion-less in the nest. After clearing the site of intruders,the male enters the nest with fins erect and buccalcavity extended. He gently pushes the female in thehind part of her body and so encourages her to startcircling around. During circling the male vibrateshis extended buccal cavity. After two to three unin-terrupted rounds, the female suddenly slows downand lay some eggs. The male too stops circling andwaits impatiently for the female to move from thesite and leave the eggs to be fertilized. One to eighttwo-millimeter long eggs are deposited at a time.As soon as the female moves forward the maleshoots over the eggs and releases its milt. The eggsare thus fertilized outside the female’s mouth. Atthe end of the spawning cycle the female will notrelease any more eggs, but the ritual goes on longafter that. Every time she moves forward the maleshoots like an arrow over the barren sand. Some-

times a male can be so excited that it shoots overthe female when she does not move fast enough.

The batch sizes vary between 30 and 80 eggs forthe two species of Enantiopus and between 10 and40 for X. ochrogenys. A male may spawn with twofemales on one day. In a short period all femalesare gravid. After three weeks the fry are releasedand are rather small. In one year they are matureand might spawn in the next breeding season.

During spawning, in captivity, the pair is fre-quently disturbed by other fish. The male spendsmuch time chasing away intruders. It is possiblethat the female gets upset by these frequent inter-ruptions and she may need a break for half an hour.Once I observed that the female continued spawn-ing with another male, after such a break. On an-other occasion I noticed that a badly disturbed pairof E. melanogenys quit spawning and that the fe-male spat out the few eggs some hours later. Thenext day she continued spawning with the samemale.

E. melanogenys, E. sp. “Kilesa” and X. ochro-genys are great residents for the Tanganyikaaquarium. We should not mix them with large orrough fish which will very likely damage the some-what fragile females. If we want to enjoy the spec-tacular view of a male in full color, we must givehim a rival. Two or more males enhance each oth-er’s activity and bring out the best colors. If onlyone male is kept, with several females, spawningtakes place but usually without the sparkling colorsof the male.

References

KONINGS, A. (1988) Tanganyika Cichlids. VerduijnCichlids, Rotterdam, Netherlands.

POLL, M. (1956) Résultats scientif. expl. hydrob.belge au lac Tanganika (1946-1947). PoissonsCichlidae, vol. III, fasc. 5B

POLL, M. (1986) Classification des cichlidae du lacTanganika. Tribus, genres et espèces. Mémoiresde la Classe des Sciences. Acad. Roy. Belg. Coll.in-8°-2e série, T. XLV Fasc. 2.

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In spring 1988, Walter Dieckhoff discovered a newLamprologine along the Tanzanian coast, near Ikola.He managed to collect five specimens alive but even-tually only two survived the strenuous trip to Europe.These two fish happened to be a male and female. In abare tank they didn’t look much different from a vari-ant of Neolamprologus brichardi, but Walter assuredme that they behave differently. He noticed that theCygnus, as I named them later, live in pairs or solitaryand not as gregariously living pairs which is knownfor N. brichardi. Walter further observed that theCygnus remains close to the rocks—a little like N.furcifer—and that they have a dark blue-brown col-oration with conspicuous blue eyes. I was fortunate toget this pair. I first placed them in a large tank withother fish in order to get them acclimatized to the typeof water and to each other. The male and female har-monized well together and I decided to put them to-gether in a small breeding tank (50 litre) which wasdecorated with rocks and a flower pot. Even at theirrelatively large size of about 9 cm the pair did well.After about two months I noticed the first fry. Therewere three in total. With Cygnus it is not the quantitybut the quality that counts. These three babies werethe most gorgeous lamprologus I had ever seen. The

At a size of about 2 cm, a juvenile Neolamprologus sp. “Cygnus” shows its best coloration.

A wildcaught male Neolamprologus sp. “Cygnus”.

Neolamprologus sp. “Cygnus”

yellow-orange fins and patches on the head were ab-solutely unique. About two weeks later the pairspawned again – only one youngster made it to thefree swimming stage. The pair kept on spawning aboutevery fortnight for more than a year. The broods, how-ever, never contained more than 16 fry. In females theyellow coloration on the head remains much longer thanin males but is lost – except for a faint tinge – at a size ofabout 5.5 cm. Tankraised Cygnus have larger nests, evenat a smaller adult size. When kept in a dimly lit tank, apair with 20 mini-butterflies is an unforgettable sight.

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Altolamprologus calvus is a small predator cichlidwhich lives in the southwestern part of the lake. Itsmaximum size is about 15 cm, but most adult indi-viduals measure between 7 and 10 cm for femalesand between 10 and 13 cm for males. The female –at the same age as the male – is considerably smallerthan the male.

Three different geographical variants are knownfor A. calvus. The holotype is from the populationwith the deepest black coloration which is found inthe southern part of the Zairean coast and inCameron Bay. The White Calvus occurs aroundCape Chaitika which is probably the most easternpopulation. Here it shares the habitat with the Yel-low race of Altolamprologus compressiceps.

In 1988 René Krüter discovered a completely yel-low race of A. calvus at Chilange Rocks. This loca-tion lies between those of the black and the whitepopulations. Since then the Yellow Calvus has be-come a valuable addition to the Tanganyikaaquarium.

The best way to keep Altolamprologus in a tankis to restrict oneself to one pair only. At a size ofabout 5 cm the Calvus can be sexed by inspectingthe vents. To be sure you should check several speci-mens to get the idea how a female and male should

A wildcaught male Altolamprologus calvus from Chilange Rocks, Zambia.

Altolamprologus calvus (Poll, 1978)

look. Females, which are much smaller than males,have a distinctly broader genital papilla than males.The larger the fish the easier it becomes to “vent”them. The chosen pair are best put together withother fish in a community aquarium. You can keepeither A. calvus or A. compressiceps in one tank butavoid having both species together.

When A. calvus is not breeding, it does not showany territoriality. Breeding starts when a femalestarts guarding an empty shell. Females have beenobserved breeding in a shell in the lake as well. Itis important to use a small shell. I have spawnedseveral pairs of A. calvus on a monthly basis usingan empty Lanistes nyassanus shell. This is the snailfrom Lake Malawi. In their natural habitat, A. calvusnormally choses a small pocket in a rock, especiallywhen the female is to big to fit in a Neothauma shell.The male should not be able to enter the shell.

During spawning, when the female sticks her eggsto the inside of the shell, the male will be over theentrance of the shell and discharges its milt. The bestsign that the pair have spawned is the observation thatthe female stays, most of the time, inside the shell.The male will be territorial now and defend the shellwith great vigor. About two weeks after spawning thefirst fry appear in the opening of the shell.

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Neolamprologus sexfasciatus occurs in the south-ern half of Lake Tanganyika. It is mainly found, inshallow water, in the intermediate biotope wheresand is present between the small heaps of rocks.N. sexfasciatus occurs solitary or in pairs. Breed-ing pairs normally have youngsters which they de-fend until they produce the next spawn. Sometimesjuveniles of a size larger than 3 cm are still guardedby the parents.

Three different geographical races are known.The holotype was collected on the Zairean coastand probably looked like the fish in the pictureabove. The first variety that was exported came fromZambia (see photo). It occurs along the entire Zam-bian coast except for the eastern part near the Tan-zanian border. The populations along the Tanzaniancoast are completely yellow and have been exportedseveral times.

The Golden Sexfasciatus, as this race is called,has been successfully bred in captivity but the BlueSexfasciatus from Zambia never.

In behavior (as far as one can judge from obser-vations in the aquarium) there is a difference be-tween these two races too. The Blue Sexfasciatusshows a much more piscivorous appetite than theGolden variant. The former cannot be kept with

A wildcaught female Neolamprologus sexfasciatus from an unknown location in Zaire.

Neolamprologus sexfasciatus from Zambia.

Neolamprologus sexfasciatus (Trewavas & Poll, 1952)

fishes smaller than 5 cm! The race from Zaire seemsto have a similar behavior to the Golden Sex-fasciatus.

It is impossible to keep more than one pair in anaquarium. In very large tanks (more than 750 liter)two or three females might do well, but usually theyquarrel among each other.

N. sexfasciatus is a cave-brooder and we there-fore have to supply the pair with some caves. Thebest way to proceed is to let the female adapt to hernew environment first. When she is acclimatizedthen the male can be introduced.

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Neolamprologus leloupi was described by Poll (in1948) from a single specimen which was collecteda little north of Moba, on the Zairean coast. Thirtyyears later, the same author described Neolampro-logus caudopunctatus, a cichlid from the Zambiancoast. This fish closely matches N. leloupi in ana-tomical features but lacks the distinct black marginin the tail fin. In 1988 Walter Dieckhoff discoveredtwo other populations of Leloupi-like cichlids onthe Tanzanian coast. One of these cannot be distin-guished from the holotype of N. leloupi. The otherone closely resembles the Zambian N. caudo-punctatus.

Pierre Brichard (1989) concludes in his monu-mental book that N. leloupi occurs in different geo-graphical populations and that N. caudopunctatusshould be regarded as a synonym for N. leloupi.

Recently Janicki imported another variant fromthe shores of Zaire. A female of this race is shownin the picture above. The trade name, under which Iacquired these pretty fish, is “Red Fin Caudo-punctatus”. I think that Brichard’s opinion will beaccepted by most taxonomists, so we should callthis little cichlid Neolamprologus leloupi.

N. leloupi lives in the intermediate habitat, whererocks lie scattered over a sandy bottom. In Zam-

A female Neolamprologus leloupi showing a defensive posture.

Neolamprologus leloupi (Poll, 1948)

bian waters it is one of the most abundant cichlidsin this biotope. Schools of hundreds of individualsare a common sight.

In the aquarium, N. leloupi behaves rather peace-fully as long as we keep just one pair. Most of themale’s aggression is directed towards conspecificmales. It can be kept with very small fish too. N.leloupi is mature at an age of about one year andmay spawn at a size of about 4.5cm. Although ithas been observed spawning in an empty Neo-thauma shell, the pair prefers something larger thanthat. If no cave is present they will start diggingone under a stone. During the process the entranceto the cave is concealed by a wall of sand (or gravel).If we provide the pair with a small flowerpot, withits bottom removed and placed upsidedown in thesand, we can chose the position of their territory.They will readily accept the pot and try to concealit by throwing sand against it. The result is a heapof sand with only the top of the pot sticking out.The size of the territory depends on the size of thefish and the population density of the aquarium, butusually it is about 30 cm in diameter. Any intruderis chased from this area but youngsters (to a size ofat least 2.5 cm) from a previous spawn are toler-ated.

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Neolamprologus mustax is not a common cichlid,either in the wild or in captivity. Its first introduc-tion, in the mid-seventies, was hampered by a wrongidentification and by the fact that a fake colorationwas applied on a picture of the Cameron Bay vari-ant, the so-called “Blue Mustax”.

The first specimens exported to Europe werecaught in Cameron Bay. The fish of this populationhave a grayish-yellow body color and yellow fins.But this race displayed mainly the rather dull graycolor when held in dealer’s tanks. Nevertheless, itwas named “Blue Petricola”. It was then still anundescribed species but some thought it could beN. petricola.

N. mustax is reported only from the southern partof the lake where it has been found in Zambianwaters. It is a small, rock dwelling cichlid with amaximum size of approximately 9 cm. Several dif-ferent populations are known. Most of them havefish with yellow fins. Some populations, like thoseat Cape Chaitika and near Mpulungu, have com-pletely yellow or orange-yellow individuals. Thename Mustax means beard which reflects the whitecoloration on the chin and lower part of the headand which is seen in all known populations.

A wildcaught Neolamprologus mustax, collected near Mpulungu, Zambia.

Neolamprologus mustax (Poll, 1978)

In its habitat N. mustax is difficult to observe asit mainly lives in the dark cracks between the rocks.It is thus also difficult to collect which makes it arather expensive Tanganyikan cichlid.

N. mustax belongs to a group of rock dwellingLamprologines in which we also find the betterknown N. leleupi, N. longior and N. cylindricus.All of these species live solitary or in pairs whenadult. They are rather intolerant of the presence ofconspecifics. It is therefore difficult to accommo-date several individuals in one aquarium. It is a goodidea to acquire just one male and a female. If thetank is large enough (about 750 liters with lots ofrockwork) both may be introduced at once but it isbetter to let the female acclimatize first.

If not properly fed or kept, N. mustax may loseits deep yellow-orange coloration. It needs a lot ofcarotene-rich food like Cyclops, Mysis or other al-gae-eating crustaceans. Besides a lot of shelter inthe rockwork it needs clean water with a high pH,preferably above 8.0. The colors of N. mustax mayfade initially but after treating it right, it will dis-play the golden yellow as brightly as in the wild.

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Greenwoodochromis christyi was observed underwater for the first time in 1988. Walter Dieckhofffound a few individuals in the Zambian part of thelake. He collected a few specimens of which only afemale survived transportation back to Europe. Oneyear later some christyi were exported by ChrisBlighnaut in Mpulungu.

Greenwoodochromis christyi belongs to a groupof Tanganyikan cichlids which have a colorationpattern consisting of a few horizontal rows of iri-descent scales. A well-known cichlid from thisgroup is Limnochromis auritus. Previously, all thesecichlids were placed in the genus Limnochromis.The species involved are L. auritus, L. abeelei, L.staneri, Greenwoodochromis christyi, G. bellcrossiand Gnathochromis permaxillaris (see next page).The difference between the species in Green-woodochromis and those that remain in Limno-chromis is that the former have more than 48 scalesin an horizontal row between the gill-cover and thetail. The species in Limnochromis have a maximum40 scales in a row. In view of the recent revision ofthe Malawi cichlids, where the genera are distin-guished mainly by their basic coloration pattern, it

A female Greenwoodochromis christyi, collected in Zambian waters.

Greenwoodochromis christyi (Trewavas, 1953)

would not surprise me if all these species wereplaced back in their original genus in future.

The two species currently in Greenwoodochromisare distinguished from each other by the size of theeye and the shape of the mouth. G. christyi has arelatively smaller eye than G. bellcrossi. The latteralso has a steeply inclined mouth, while that of G.christyi is only moderately inclined.

Greenwoodochromis christyi has been observedin the intermediate habitat at somewhat deeper lev-els. René Krüter found most individuals at a depthof 25 meters and deeper. Sometimes a pair was ob-served while guarding their offspring. Spawning hasnot been observed but it is probably similar to thatof L. auritus. This means that G. christyi is amouthbrooder where both female and male take careof their offspring.

Greenwoodochromis christyi has proven to be arather difficult cichlid to keep in an aquarium. Asingle specimen can be kept, without problems, withother species of similar size but a “pair” need a verylarge tank. All their aggression is directed towardsconspecifics. The solution to this problem could beto let the female adapt to the tank first.

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One of the most remarkable cichlids of Lake Tan-ganyika is Gnathochromis permaxillaris. Its mostinteresting features are the very large mouth whichopens in a peculiar way (see photo on the backcover)and the shape of the upper lip. It is a rather largecichlid which is known to grow to a size over 18’cm.

Gnathochromis permaxillaris lives over muddysubstrates and is usually found deeper than 40 me-ters. Although the enormous mouth may suggestotherwise, mostly very small organisms constitutethe staple food of G. permaxillaris. These tiny in-vertebrates live close to the bottom, sometimes inthe upper layer of the mud. The wide gape ofpermaxillaris spreads the inflow of water, going intothe mouth when the gill covers are extended, over arelatively large area. This weakens the flow so thatonly small and light particles are carried in. Thelarge cichlid has to “vacuum clean” the muddy bot-tom continuously in order to obtain enough mate-rial to live on.

Gnathochromis permaxillaris is probably the most interesting cichlid from Lake Tanganyika.

Gnathochromis permaxillaris (David, 1936)

The same feeding technique is observed in theaquarium where G. permaxillaris feeds predomi-nantly from the bottom. It takes a while before itadapts to picking up food in mid-water.

Gnathochromis permaxillaris is now regularlyexported from Zambia albeit in very small numbers.It seems to be a very fragile fish to collect but whenacclimatized to the aquarium it proves to be a hardyand undemanding fish. A very welcome factor –completely in contrast to Greenwoodochromischristyi – is its remarkably peaceful attitude towardsconspecifics.

The sexes can be identified by looking at the ven-tral apertures; the genital opening of the female isnoticeably larger than that of the male. Unfortu-nately, there are usually not enough specimens tocompare and two males or two females might hidein the same cave without showing any hostility to-wards each other.

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The cichlids of the genus Trematocara belong to thedeepest living fishes of Lake Tanganyika. Representa-tives of this genus were frequently captured at a depthof 200 meters. It is assumed that these cichlids mi-grate freely between the very deep layers and the sur-face layers of the lake. Especially at night they moveto shallower water. This means that a special construc-tion of the swimbladder allows the fish to compen-sate for the enormous difference in pressure. Duringthe daytime Trematocara lives in the depths probablytogether with its food. It is known that a small crusta-cean, a kind of shrimp, undergoes a diurnal verticalmigration as well. Masses of this shrimp are found inshallow water at night.

Evolution has permitted Trematocara nigrifrons tofollow these crustaceans during their daily migrations.This means that these cichlids live in permanent dark-ness. They have well developed adaptations to thiscircumstance; T. nigrifrons possesses a very large eyeand an extremely sensitive lateral organ on the headwith which it can register minute movements in thewater. The sensory pore system, which is present onthe head of other cichlids as well, is greatly enlargedon the head of Trematocara. The organ, which con-sists of pressure sensitive tubules, is visible as small“holes” below the eye. In the aquarium, I noticed that

A male Trematocara nigrifrons collected in Zambian waters.

Trematocara nigrifrons Boulenger, 1906

food is only accepted (greedily) when it moves. Fro-zen food must be stirred by other fishes before T.nigrifrons eats it. I have seen it hovering above thefood on the sand, in a manner similar to Aulonocara,and waiting for a movement to occur.

Roger Bills, who is a researcher and collecter ofZambian fishes, managed to catch and ship ten speci-mens of T. nigrifrons. Unfortunately the fish provedto be very sensitive and, after one year, only three sur-vived. Surprisingly I succeeded in breeding them. Themale is considerably smaller than the female and, whenexcited, shows black and white edges to the fins and ablack throat. Regrettably I did not observe the spawn-ing; I first knew of it when I noticed the female with amouthful of pale yellow eggs. After three weeks thefry were about 8 mm in size, when the female spatthem out because of a stress caused by netting her outof the tank. The approximately 40 fry immediatelyate Artemia and now, three months later, they are about4 cm long and am I the proud owner of a school of T.nigrifrons.

T. nigrifrons is not a colorful addition to anyaquarium but for a true Tanganyika fancier, who usu-ally looks further than colors alone, it is a very inter-esting and elegant species.

René Krüter

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The specimens depicted in the photograph above areonly tentatively identified as Microdontochromistenuidentatus. The holotype of this species shows, al-beit in preserved condition, a horizontal, mid-lateralrow of black spots. In some preserved specimens thesespots appear as vertical bars. The cichlids in the pic-ture above show only such pattern when they are ex-cited. On the other hand, their morphology and theirtypical mouth strongly suggest that these fish areconspecific with M. tenuidentatus.

The holotype, together with about 50 other speci-mens, was collected in Zaïre near Vua. The fish in thepicture were collected in Zambia in Cameron Bay,which is not far from Vua. Sporadically, Fishes ofBurundi has exported some specimens which alsomatched M. tenuidentatus in coloration even when theywere not excited. These were doubtless collected inBurundi waters.

It is known that Cameron Bay harbors some varie-ties (or even distinct species) which are not found atany other place in the lake, for example Petrochromistrewavasae, the so-called Tiger Nasutus or the NdoleBay variety of Xenotilapia ochrogenys. It is thereforepossible that the nondescript cichlid, when not excited,is a geographical variant of M. tenuidentatus which is

Two wildcaught specimens of Microdontochromis tenuidentatus of unknown sex.

Microdontochromis tenuidentatus (Poll, 1951)

only to be found in Cameron Bay.Microdontochromis tenuidentatus is a maternal

mouthbrooder. It has been bred in captivity by VolkerPuttberg, Dinslaken, Germany. He noticed that the eggswere rather large and that a female broods only a feweggs at the time, although about 20 are deposited dur-ing a spawning. The male does not seem to make anest or defend a territory. The eggs are brooded forabout four weeks before the fry are released. Duringthis period the female continues feeding like she hasnothing in her mouth not to be swallowed.

In the lake M. tenuidentatus lives in large schoolsin very shallow water. Most of them are found in thevicinity of aquatic weeds. The mouth is quiteprotrusible and the upper arm of the upper jaw bulgesthe skin of the snout. The head is laterally compressed,which is an important characteristic of this species. Itseems that M. tenuidentatus feeds from the planktonin the open water as well as from invertebrates whichlive among the weeds.

The teeth in the outer jaws are minute and look ratherfragile. The lower pharyngeal bone is very thin andslenderly shaped. The teeth on this bone are minuteand pointed. The diet of M. tenuidentatus must con-sists of very soft invertebrates and or tiny organisms.

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Cichlids from the genus Petrochromis are easilyrecognized by the structure of the mouth. The nu-merous teeth in the large jaws are permanently vis-ible in most species. Petrochromis orthognathus isthe only known species in the genus in which theteeth are less numerous and where the teeth are in-visible when the mouth is closed. A cichlid withthe trade name “Petrochromis Tricolor” was ex-ported from Zambia. The specimen which I was ableto have a closer look at showed just a few rows ofteeth in the mouth. It could close the mouth com-pletely, “hiding” all teeth. The coloration patternclosely matches that of P. orthognathus, i.e. a brownupper half of the body and a lighter lower half.

Petrochromis polyodon occurs in several differ-ently colored populations of which at least five havebeen exported. The cichlid we have previouslycalled Petrochromis famula is the polyodon racefrom Kigoma. Petrochromis polyodon is easily rec-ognized by the short lower jaw and the generallybrown coloration of the females. Petrochromismacrognathus also has a short lower jaw but itsmouth is underslung.

The “Red Texas Petrochromis” is closely related tothe “Texas Petrochromis” from the Tanzanian coast,but is collected on the Zairean side of the lake.

A Zairean race of Petrochromis polyodon which is named “Kaiser Petrochromis” in the trade.

Above: The “Red Texas Petrochromis”; below:

Petrochromis, three new variantsAd Konings

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Many different populations of Tropheus moorii areknown and many more will be found in the future.The genus Tropheus consists of at least six speciesof which one is scientifically undescribed. This spe-cies, named T. sp. “Black” or—previously—T. sp.aff. brichardi, lives in the northern half of the lake,whereas T. moorii is restricted to the southern half.Although both species have been regarded asconspecific by several authors, Walter Dieckhoffdiscovered a locality where both species livesympatrically (Kibwesa, Tanzania).

A wildcaught male Tropheus moorii, probably from Vua, Zaïre.

Tropheus moorii from an unknown location in Zaïre.

Tropheus moorii Boulenger, 1898Ad Konings

In Zambia and along the lower half of the Zaïreancoast the populations of T. moorii can be split intotwo different groups. The holotype of T. moorii wascollected near Mpulungu and belongs to the groupof light-colored races which are further character-ized by a colored region on the center of the body.In Cameron Bay and north of it, populations of thesecond group are found. This group consists of dark-colored races which also have a colored cheek. Thereis no location known where individuals of both groupslive together which would have indicated that we aredealing with two different species.

Although there is a rather large difference, in col-oration, between the two groups (see photographs),they probably belong to the same species. The twogroups therefore represent two main geographicalvariants. The Kasaba Bay in Zambia might haveseparated these two groups for a long period of timeand they might thus have developed, independentlyfrom each other, into the present races. The north-ern borderline between the two groups is not knownbut a dark-colored variant has allegedly been col-lected near Vua in Zaïre. Further north into Zaïreraces of the light-colored group have been found.One of them is shown in the smaller picture.

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Cyathopharynx furcifer is a rather common andspectacularly colored cichlid from Lake Tangan-yika. Males attain a total length of about 18 cmmaximum; females, usually, are considerablysmaller. Cyathopharynx furcifer lives mainly oversandy areas in shallow water. However, such sandybays must be close to rocky shores. The reason whythey prefer the rocks nearby is unknown becausethey predominantly feed from the sandy substrate.

The food consists of diatoms which are collectedfrom the upper layer of the sediment on the sand oron the rocks. Large quantities of material arescooped up and processed through the long intesti-nal tract. While feeding, the faeces can be seengrowing from the vent. A lot of the material swal-lowed is indigestible. This results in an easy pas-sage of the food through the guts. We must keepthis fact in mind when maintaining this cichlid inthe aquarium. Cyathopharynx furcifer needs a lotof food but only a little meat.

In the lake, males congregate into large breedingcolonies. Each male makes its own nest which is akind of sand-castle with a diameter of approximately50 cm. The nests are about 30 cm high. The differ-ent territories can be just two meters apart. Breed-

This race of Cyathopharynx furcifer lives on the Ruziba reef in the northeastern part of the lake.

Cyathopharynx furcifer (Boulenger,1898)

ing colonies of C. furcifer are a spectacular sight;sometimes more than 100 colorful males are foundtogether defending their nests. The females remainin large schools, normally close to the rocky coast.

Because C. furcifer seems to be dependent onareas with rocks, which are not present along theentire coast uninterruptedly, it has developed sev-eral geographical races. The Ruziba race, which isshown in the photo, was exported from Burundi afew years back.

When C. furcifer is collected the fishermen catchthe most beautiful males. These males keep theircolors for just a few weeks. Rarely do they regaintheir full palette of colors in captivity, even thoughthey might spawn and produce offspring. Tankraisedmales, however, show their best colors when stillsmall and usually keep them until something largeor rough disturbs them in the aquarium. In theaquarium they need a large area with plain sandwhere they can build their nest. Only in very largetanks (over 750 liters) can we perhaps have morethan one male in color. The presence of a second,smaller, male usually results in the most dominantmale retaining his colors.

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Xenotilapia flavipinnis is a common, sanddwellingcichlid which is found all round the lake. It is foundpredominantly over sand at depths between 1 and30 meters. X. flavipinnis lives in schools. The larg-est schools are found in the very shallow water.Foraging schools may number over a hundred indi-viduals. Like all other members of Xenotilapia itfeeds by sifting the sand. The food consists mainlyof soft-bodied invertebrates, like insect larvae.

It seems that X. flavipinnis has traditional forag-ing and breeding grounds and that it is thus restrictedto certain areas. This has probably led to the exist-ence of geographical races. Three of these raceshave been exported to date. The holotype belongsto the northern race of Magara, Burundi. The yel-low variant from Nyanza Lac, Burundi, is shown inthe picture. From Zambia a variant is frequentlyexported which shows a few rows of yellow scaleson the flank.

Xenotilapia flavipinnis breeds in colonies but itis not a lek-breeder. Pairs separate from the school– the school splits up into many breeding pairs –and establish a rather large territory. Even thoughwe speak of a breeding colony the pairs are usuallymore than two meters apart. For a small cichlid –the maximum size of X. flavipinnis is about 10 cm

The yellow Xenotilapia flavipinnis from Nyanza Lac.

Xenotilapia flavipinnis Poll, 1985

– this is a large territory. In order to avoid territo-rial disputes in the aquarium it is therefore best keptas a pair.

Spawning occurs at any site in the territory. Thesite might change during one spawning. There isno visible nest constructed but it seems that the pairchooses a slight dip in the sand. A few days beforethe actual spawning, the male courts the female bypositioning himself with all fins erect and the buc-cal cavity extended. The female answers in a simi-lar way. Without much introductory behavior thefemale starts laying the eggs which are fertilizedby the male while they are still on the sand. Afterthe female has taken all the eggs in her mouth shestays with the male in the territory. After about 9days the larvae are spat on the sand, in front of themale who picks them up immediately. The malecarries the larvae for another 5 to 6 days before hereleases the fry in the territory. The fry are tendedby both parents – mostly by the male – for anotherweek or two. A further spawning usually occurswithin a month in the same territory.

Xenotilapia flavipinnis is a rather peaceful cichlidwhich is best kept with non-predatory cichlids inan aquarium with fine sand on the bottom.

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Xenotilapia papilio was described from a popula-tion 40 kilometers south of Moba on the Zaireancoast. The two individuals shown on this page verylikely belong to the same species and are thus geo-graphical variants. X. papilio is a very smallsanddwelling cichlid. Its maximum size lies around8 cm. This very pretty species is mainly character-ized by the underslung and tiny mouth. The teethare bicuspid, which excludes them from the genusAsprotilapia which is characterized by tricuspidteeth. However, X. papilio closely resembles A.

Several geographical variants of Xenotilapia papilio occur in the southeastern part of Lake Tanganyika. This race isknown as the “Sunflower Xenotilapia”.

Xenotilapia papilio. This variant is known as “Chituta BayXenotilapia”.

Xenotilapia papilio Büscher, 1990

leptura and is also found sympatrically with thelatter.

Xenotilapia papilio lives in shallow water but atsome locations it may penetrate into depths of morethen 30 meters. In the deeper areas is is commonlyfound in intermediate habitats, where sand and rocksare both present. In very shallow areas it occurs overpure sand. Large schools have never been observed.Usually pairs or small groups are sighted. Like X.flavipinnis it pairs off and defends a territory. Ob-servations of aquarium populations show that oncea pair is formed they stay together for several con-secutive spawns. Xenotilapia papilio is best keptas a pair in a community tank as long as there areno predatory species accompanying them. They re-gard similar looking species, such as X. flavipinnis,as competitors which elicit frequent quarrels.

Spawning in X. papilio is preceded by fin-dis-plays by both male and female. One or two dayslater the female deposits the eggs on a site some-where in the territory. These are fertilized by themale while they are still on the sand. The femaleinitially broods the eggs. The male takes the larvaeafter about 8 to 10 days. After less than a week thefry are released.

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Xenotilapia sp. “Katete” looks similar to the recentlydescribed X. papilio but definitely is another species.It has a shorter snout, a larger eye, a differently shapeddorsal fin and a shallower caudal peduncle. The maxi-mum length of X. sp. “Katete” lies around 7 cm, alittle bit smaller than X. papilio. In contrast to latterspecies, the dorsal fin of X. sp. “Katete” is entirelyorange in color. The spines of the anterior part arelonger than those of the rest of the fin. The body has afaint blue hue caused by a few rows of blue-reflectingscales. Personally, I find this the most attractive spe-cies among the small, rock-dwelling Xenotilapia.

The village of Katete lies near the border betweenZambia and Zaïre. In October 1988 we were lookingthere for a good spot to collect Cyphotilapia frontosa.While I was diving at 35 to 40 meters, I noticed X. sp.“Katete” but at first glance I thought it was X.flavipinnis. Realizing that that species is not commonlyfound at these depths I looked closer and recognizedit from a picture from Horst Walter Dieckhoff in AdKonings’ book.

Xenotilapia sp. “Katete” inhabits the intermediate zonebetween sand and rocks, usually on a steeply sloping partof the coast. C. frontosa and Neolamprologus buescheriwere among the other fishes of its habitat, which would

Xenotilapia sp. “Katete” lives at the deeper levels of the rocky habitat.

Xenotilapia sp. “Katete”

be a bad combination for the aquarium.With a lot of effort we could only collect a small

number of these fragile cichlids of which just five sur-vived the transport to my tank. They were three fe-males and two males. There are no clear morphologi-cal differences between the sexes. Unfortunately, X.sp. “Katete” is very selective when it has to form pairs.I was unlucky that none of the five specimens reallyliked each other. As long as there had not been a spawn-ing the pair seemed to be doing well; the other three“Katetes” were then “expelled” from the 1000 literaquarium. These three were housed in a similar sizedtank of my friend Willem Bastinck.

Xenotilapia sp. “Katete” is a biparental mouth-brooder, this means that the larvae are transferred tothe male after the female has brooded them for about10 days. The problem with my pair started when themale had taken the larvae in his mouth, because thenthe female started to demolish the male, who reactedby spitting out the larvae. Fortunately, Willem hadmore success and was able to save some of the larvaeafter the male had spat them out. Later, my pair also pro-duced some fry which I was able to save from the fe-male’s aggression. Now, after three years, the aquaristicfuture of these beautiful cichlids looks a bit brighter.

René Krüter

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Cyprichromis sp. “Leptosoma Jumbo”

Cyprichromis sp. “Leptosoma Jumbo”, a morph of the Kampemba race.

Cyprichromis sp. “Leptosoma Jumbo”, a yellow-tailed morph from an unknown location in Zaïre.

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The small cichlids from the open water in Lake Tan-ganyika belong mainly to the genus Cyprichromis.There are only two species described in this genusbut at least two others are known. The main differ-ence between these species lies in the color patternsof the males. They have gorgeous colors which areclearly seen in their natural environment. Cypri-chromis is spread over the entire lake but only C.microlepidotus occurs in the northern part, north ofNyanza Lac. From observations in Zambian waters(Walter Dieckhoff; René Krüter, pers. comm.) it isknown that several species of Cyprichromis maylive in direct contact with each other.

Cyprichromis microlepidotus, which has a lake-wide distribution, is characterized by small scales.These are smaller than those from any other knownspecies of the genus. It is also larger than C.leptosoma, which is a small and slenderly builtcichlid. The latter species differs from C. sp.“Leptosoma Jumbo” by its shallow body and smalleradult size only. C. sp. “Leptosoma Jumbo” can growto a size, at least under aquarium conditions, of ap-proximately 12 cm whereas C. leptosoma rarelyexceeds a length of 9 cm. The fourth species, C. sp.“Leptosoma Zebra”, is recognized by its yellowishcoloration – its lack of conspicuous colors – and bythe broad vertical bars which are not permanentlyshown. C. leptosoma (Blue Flash, Isanga race), C.microlepidotus, (Yellow Zambia race), C. sp.“Leptosoma Jumbo” (Black Fin), C. sp. “LeptosomaZebra” and Paracyprichromis nigripinnis (BlueNeon) have all been seen together in a small area inChitutu Bay.

C. sp. “Leptosoma Jumbo” is a pleasant aquariumfish which normally shows its full colors. It is im-possible to keep this species in pairs. Males activelydefend a territory and need the presence of at least

Cyprichromis sp. “Leptosoma Jumbo”, a morph from theKampemba population.

C. sp. “Leptosoma Jumbo”, a male of the yellow-tailedmorph from the Zaïrean race.

four other males to minimize the damage done tothe weaker one. Males behave rather peacefully to-wards the females. In three breeding colonies, thatI have kept over several years, I have never found afemale killed by an aggressive male. Depending onthe number of males in the aquarium, they will stakeout their territories about 30 to 60 cm apart. Themales stay in mid-water and relate the size of theirpremises to the distance from neighboring males.Females are continuously courted and attracted tothe territory. When a female reacts to the male’sinvitation, the male bends its body and all its fins,except the ventrals, away from the female. The ven-tral fins, which have a large yellow spot, are heldto the outside, in front of the female’s mouth. In-stinctively, the female snaps at the male’s ventralfins. Since spawning takes place in mid-water, thisaction might indicate to the female the spot whereshe should deposit her eggs. The color of the spotin the ventral fin does not exactly match that of theegg. I have also observed males snapping at the pre-sented fins of a more dominant male.

When a female is ready to spawn she remainsinside the male’s territory. Initially she will snap atthe male’s ventrals several times before she startsdepositing the eggs. He might discharge his miltbefore the eggs are laid. At a certain moment themale positions himself over the female, all fins ex-tended, and pushes her gently, with his fully openedmouth, on her head. The female releases one or afew eggs and immediately backs up to retrieve it.The sequence of release and backing up may be re-peated several times before the female is led back,by the male, to the center of the territory. Then fol-low a number of “ventral-snappings” before the fe-male discharges the next batch of eggs. The incu-bation period lasts about three weeks.

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Of all the new imports of the last years, the “BlueNeon Cyprichromis” is the favorite of most Tan-ganyika cichlidists. It is a small and peaceful cichlidwith a gorgeous coloration. The maximum size ofParacyprichromis nigripinnis is about 10 cm foraquarium specimens. Its shape closely resemblesthat of P. brieni, the other species of the genus. Thedifference lies mostly in their preferred habitatdepth, although P. nigripinnis has a noticably largereye (relatively seen). The latter is also found indeeper regions of the rocky coast than P. brieni,which lives in the upper 10 meters. The “Blue NeonCyprichromis” is caught at a depth between 20 and30 meters. In Chituta Bay it lives in large caves orin close vicinity of large boulders.

Paracyprichromis nigripinnis has a lake-widedistribution but not all populations have such anattractive pattern as the race shown in the photo-graph. René Krüter (pers. comm.) found two dif-ferent populations in Chituta Bay. The differencebetween the two is the color of the dorsal fin, whichis more red in the other race. Walter Dieckhoff founda race in Kigoma Bay which does not show the“neon” stripes.

Paracyprichromis nigripinnis and P. brieni werepreviously included in the genus Cyprichromis but

Paracyprichromis nigripinnis from Chituta Bay is known as “Blue Neon Cyprichromis”.

Paracyprichromis nigripinnis (Boulenger, 1901)

because of the difference in the number and arrange-ment of the vertebrae they were placed in a newgenus. Although the splitting-up might seem a mat-ter of opinion, P. nigripinnis and P. brieni are prob-ably not at all related to the species currently inCyprichromis. In my opinion, they just show a re-markable case of parallel evolution, whereby spe-cies of two different ancestral origins have devel-oped into similar looking species.

The most important distinction between these twogenera, however, lies in their spawning behavior.The eggs of Cyprichromis are fertilized inside thefemale’s mouth (see page 27) but those of Para-cyprichromis are not.

Male P. nigripinnis defend a territory alongsidea large rock or inside a cave. Females are not at-tracted to the nest by fin display but are seques-tered from the school and then led to the male’sterritory. The female swims head-down, close to thesubstrate, and discharges the eggs. As the eggs passby her head they are collected by the female. Mean-while, the male stays above or beside the femaleand discharges – visible clouds of sperm are occa-sionally seen – its milt continuously. He fans themilt towards the falling eggs which are thus ferti-lized before the female picks them up.

Ad Konings

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Of all the different biotopes Lake Malawi has tooffer, the rocky regions have brought forth the great-est variety of colorful aquarium fishes. The muchless explored sandy areas may still hide many beau-tiful species. Due to the openness of the sandy habi-tats it is difficult, and sometimes impossible, tocollect some specimens. Since on the sand there isno place to hide, almost all species living there havea light, silvery-yellow coloration which camou-flages them. Furthermore, the sanddwellers oftenlive in schools which gives them a protection against

Placidochromis phenochilus is a very conspicuous cichlid from the sand.

MALAWIANCICHLIDSThe blue sanddwellers

predators.In contrast to most sanddwellers, there is a group

of conspicuously colored cichlids which usually livesolitarily or in very small groups. This group con-sists of at least five different species. These haveno ancestor in common and do not belong to thesame genus. However, they have an important andpeculiar part of their feeding behavior in common(see later).

The five conspicuously colored sanddwellers areCyrtocara moorii, Protomelas annectens, Otophar-

Ad Konings

30

Placidochromis electra with a digging Taeniolethrinopspreaorbitalis.

The distribution of Placidochromis electra is not restricted to Likoma Island. This specimen was pictured near Fort

ynx selenurus, Placidochromis electra , andPlacidochromis phenochilus. The first three specieshave a lake-wide distribution. P. electra occursaround Likoma Island and probably all along theeastern shores, down to Fort Maguire. P. pheno-chilus, which seems to be closely related to P.electra, has been observed only in the northwest-ern part of the lake. I found it near Mdoka, Chesese,Chirwa Island, and between Selewa and Kasinda.

P. phenochilus is characterized by its white lipsand blue body. Even juveniles of 6 cm length havethe typical dark blue coloration. Its inclusion inPlacidochromis is based on its close resemblanceto P. electra. Of the about 25 individuals I have seenat several locations, not a single one showed itsbasic coloration pattern. The vertical barring, whichis faintly present in most individuals, is not a diag-nostic feature since species from several other gen-era show vertical bars – especially courting males– as well.

The five species of the group have a particularfeeding strategy. They are attracted to stirred-upmaterial and usually follow a large sand-sifting spe-cies like Taeniolethrinops preaorbitalis. The foodof most sanddwelling species consists of inverte-brates which live and hide in the sand. There are

several ways to obtain these. Lethrinops-like spe-cies dig in the sand and sift it through the gills.Species of the genus Aulonocara detect their preyby “listening to the sand”. Other species blow awaythe upper layer in order to reveal the hiding inver-tebrates. The five blue sanddwellers let other spe-cies do the work. T. preaorbitalis is a large cichlidand its continuous digging efforts create a lot ofstirred-up material. Although it seems that the bluesanddwellers select their food from the clouds ofmaterial spilled through the gills of the large sand-sifter, they are actually more interested in search-ing the ploughed sand for exposed invertebrates.

31

A male Protomelas annectens is rarely seen in its full blue breeding coloration. Photo taken near Otter Point.

Protomelas annectens is the most frequently seen follower of Taeniolethrinops preaorbitalis (photo taken near Fort

32

Otopharynx selenurus from Senga Bay

These crawl back into the sand as soon as possible.The intriguing question is, why are these

sanddwelling species so conspicuously colored?Especially P. phenochilus, O. selenurus and to asomewhat lesser extent C. moorii, always show adistinct blue coloration. These three species seemto be mostly dependent on a large, sand-sifting host.The other two species are also found foraging ontheir own. It is therefore possible that their conspicu-ous colors might warn off other followers – espe-cially other species – from joining their host. Insandy regions, cichlids commonly occur in groupsor schools, often of mixed species and it would thusbe normal for an individual to join such a group.The amount of food revealed by the ploughing ac-tivities of a large digger may not be sufficient formore than one adult follower. If the follower ishighly dependent on the host, it may signal its po-sition to the other species by taking on a territorialcoloration. In particular individuals of the samespecies are wary of approaching an “occupied” host.

C. moorii and O. selenurus occur in very shal-low water and are rarely found below depths of 10meters. P. phenochilus and P. electra are usuallyobserved in water deeper than 15 meters whereasP. annectens is found at any level.

The differences not only in the basic melaninpatterns but also in breeding behavior point to dif-ferent ancestral origins for these sanddwellingcichlids. The breeding behavior of C. moorii hasbeen observed several times by different aquarists.They all noticed that as soon as the female had de-posited some eggs they were fertilized by the malewhile they were still on the sand. The same sequenceis also employed by P. annectens. Neither of thetwo species build a nest or any structure to indicatethe spawning site. Although I have never observed

any of these blue sanddwellers spawning in theirnatural environment, the courting behavior of themales indicates that spawning can take place at anysite. Mouthbrooding and fry-guarding C. moorii andP. annectens have been observed on the open sand.

P. electra is kept and bred by many hobbyists.Spawning in the aquarium has revealed that the eggsare fertilized inside the female’s mouth, as is thecase in most other Malawian cichlids. P. pheno-chilus has yet not been exported as an aquarium fish.Because P. phenochilus resembles P. electra to agreat extent it may have a similar breeding behavior,i.e. that the eggs are fertilized inside the female’smouth.

O. selenurus is infrequently exported as Haplo-chromis Nussae. All specimens are collected inSenga Bay. It has been bred in captivity (Baasch,pers. comm.) but details of the spawning sequenceare not known yet. The juveniles show the genus-specific blotch pattern until they have reached alength of approximately 7 cm. O. selenurus is a veryattractive and peaceful cichlid and makes a valu-able addition to a Malawi community tank.

References

ECCLES, D.H. & TREWAVAS, E. (1989) Malawiancichlid fishes. The classification of someHaplochromine genera. Lake Fish Movies,Herten, Germany.

DRUMMOND, B. (1976) How I keep.....Haplochromisannectens. Bunt. Bull. (Am. Cichl. Ass.), Dec.,pp. 29-30.

KOCHER, T.D. & MCKAYE, K.R. (1983) Defense ofheterospecific cichlids by Cyrtocara moorii inLake Malawi, Africa. Copeia (2), pp. 544-547.

KONINGS, A. (1989) Malawi cichlids in their natu-ral habitat. Verduijn Cichlids, Rotterdam, Neth-erlands.

SEIGARS, D. & BERARDO, T. (1979) Spawning theDeep-Water Haplochromis. TFH, Vol. 27; April,pp 4-12.

33

In May 1989, Walter Dieckhoff and I visited thenorthern region of Lake Malawi, north of Chilumbafor the first time. Our trip took us to the SongweRiver which is the border between Malawi and Tan-zania. It took us two days and several dives to con-clude that there are no rocky reefs, coasts, or isletsbetween the border and Ngara. The entire coast,here, consists of sandy and swampy beaches al-though small rocks may lie scattered on the beachand suggest that the area is rocky.

At Ngara, the most northerly rocky area inMalawian waters, an intermediate type of habitatsupports a number of rock dwelling cichlids includ-ing Aulonocara stuartgranti, the race with the or-ange patch on the body. While diving there for thefirst time I noticed some elongated sand-coloredcichlids which moved in small groups through thehabitat. At first glance they seemed to be of thegenus Aulonocara, a species of the sand dwellinggroup. They were rather shy and I could take onlytwo photographs. At that time I did not see any in-dividual with male coloration, which could be a fur-ther indication that it belonged to the sand dwell-ing group of Aulonocara. Of course I was eager tohave some specimens for a closer examination andasked Saulos Mwale, our boat leader and an ex-

A male Aulonocara auditor patrolling its territory.

Aulonocara auditor (Trewavas, 1935)

tremely skilled fisherman, to catch a few. Saulosneeds only two words to describe the desired fishand he will catch it for you, usually much bettercolored specimens than the ones you had seen. Un-fortunately, he could not find individuals with breed-ing coloration which plainly meant they were notthere. However, two specimens were preserved andthey later turned out to be A. auditor.

At the next location where we dived, at Mdoka, Ifound the same species again, but again no breed-ing males. Further south, none of us saw A. auditoragain.

Gary Kratochvil and I went back to the sameplaces in December 1990, where I found A. auditorin breeding coloration. In contrast to the few indi-viduals I had seen in May 1989, we now observedhundreds of them in large schools. Males in fullbreeding coloration, however, were rare. Most in-dividuals of this species of Aulonocara were foundbelow 15 meters of depth. The few territorial maleswere found at about 20 m, at the deepest part of thecoast where rocks were still present.

The territorial behavior of the males resemblesclosely that of the so-called Chitande Type Aulono-cara; A. auditor might therefore belong to that groupof the genus as well. Males in breeding coloration

Ad Konings

34

Aulonocara ethelwynnae is found sympatrically with Aulonocara auditor.

Aulonocara auditor occurs in schools over sand near rocks. Aulonocara stuartgranti photographed at Mdoka.

“defend” their territories with little effort. Otherspecies and females are tolerated at all times withinthe boundaries of the territory. Competing males are“chased” by raising the dorsal fin and obstructingtheir way. Under natural circumstances, I have neverobserved fights, neither in A. auditor nor in ChitandeType Aulonocara. In the aquarium, however, fightsmay occur. Territorial males live rather far apart inthe wild and remain very static.

The territory of a male A. auditor usually lies onthe sand between some stones or rocks. They were

found only at the edge of the rocky habitat. Femalesoccur in large schools and forage over the sand.Within such schools males with faint breedingcolors were observed. These males, sometimesshowed intolerance towards each other and “chased”each other from the feeding site. No territories weredefended within the schools although the school andits members remained rather static.

Other species of Aulonocara were found in theschools together with A. auditor. Besides two dif-ferent sand dwelling species, A. rostratum and A.

35

Aulonocara sp. “Chitande Type Kande”.

sp. “Jumbo Blue”, foraging individuals of A.ethelwynnae (Northern or Chitande Aulonocara)were also seen.

Ngara and Mdoka are the only two locationswhere two different species of the Chitande TypeAulonocara group are sympatric. Notwithstandingthe observation that A. auditor has a similarbehavior and habitat preference, it is morphologi-cally clearly different from A. ethelwynnae. Thesnout of A. auditor is considerably longer than thatof all other known species of the Chitande TypeAulonocara. In all the other species the snout pro-file is rounded whereas it is straight in A. auditor.This might mean that A. auditor is able to poke itssnout deeper in the substrate than A. ethelwynnaeand thus feed from a different population of inver-tebrates. Personally I think that food is abundant intheir environment and that those two species arenot competing with each other for food. The struc-ture of the teeth in the jaws looks similar in allChitande Type Aulonocara; A. auditor is no excep-tion. The four to six rows of teeth stand in a rela-tively wide band anteriorly and become a single rowon the sides of the jaw. The flat lower jaw, with itsteeth, is somewhat reminiscent of that of severalLethrinops species. A. auditor may be more of a

sand dwelling species than any other Chitande TypeAulonocara. Territorial males of the other speciesin this group are found throughout the year; theydon’t seem to have a particular breeding season. Oneexception might be A. sp. “Chitande Type Kande”breeding males of which were rare in June 1989. InDecember 1990, this species was abundantly presentaround Kande Island and many territorial malescould be observed.

At Kande Island, A. sp. “Chitande Type Kande”shares the habitat with several other species of thegenus. Two of them, A. kandeensis and A. steveni,are found in its immediate vicinity. It might thus bepossible that A. auditor and A. sp. “Chitande TypeKande” breed mainly at a time when other speciesof Aulonocara decrease their breeding activities.These two species might be unsuccessful in obtain-ing a territory when the other species are still at thepeak of their breeding season.

References

TREWAVAS, E. (1935) A synopsis of the cichlidfishes of Lake Nyasa. Ann. & Mag. N. Hist. Ser. 10.Vol XVI.

36

Abstract

The dark-bellied piscivorous cichlid of Lake Ma-lawi is Tyrannochromis macrostoma (Regan), as it wasidentified by Lewis et al. (1986: 39), who placed itprovisionally in the genus Cyrtocara. The darker lowerhalf of the head and body is usual for adults of thisspecies in the lake, but is masked by the blue colourof breeding males and may be lost in unfavourablecircumstances. It is thus rarely seen in captive or pre-served fishes.

The name T. nigriventer Eccles belongs to a differ-ent species, related to Nimbochromis fuscotaeniatus.

The names T. maculiceps (Ahl) and T. polyodon(Trewavas) may be synonyms of T. macrostoma butthis is not yet proved.

Introduction

Our friend Ad Konings, to whom we owe so muchfor his books on the home life of the cichlids ofLakes Tanganyika and Malawi, has contributed tothe solution of the problem of the specific identityof the predator known as “Dark-belly” or “Schwarz-bauch”. But in his article (Cichlidae (BCA) Vol. 11,no. 3) he has made some conflicting statements that

A beautiful male Tyrannochromis macrostoma in breeding coloration. Photo by Stuart Grant.

The genus Tyrannochromis Eccles & Trewavas, 1989

Dr. Ethelwynn Trewavas

need clarification.Konings writes (Cichlidae 11 (3): p. 73) “The so-

called Black-belly is thus correctly named T.macrostoma”. But two paragraphs later “the speciescurrently known as T. macrostoma should be correctlynamed T. nigriventer”.

These are apparently mutually contradictory state-ments. In two publications that may be considered“current”, namely Lewis et al. (1986; p. 39) and Ec-cles & Trewavas (1989, p. 97. fig. 43), the namemacrostoma is correctly used for the species to whichRegan gave that name in 1922. Konings now tells methat by “currently” he meant “currently in theaquaristic trade”. Also in Konings’ books, 1989, 1990,macrostoma has been incorrectly used for nigriventer.This is important for us to know because all the eco-logical information we have about T. nigriventer isfound in Konings’ books under the name macrostoma.While the true macrostoma is there treated under thename T. maculiceps, which may be a synonym of T.macrostoma, or at least the name of a closely relatedspecies or subspecies.

To explain how this confusion came about let merelate the following, which also shows the value ofcooperation between field naturalists, aquarists andtaxonomists.

37

A male Tyrannochromis nigriventer in breeding coloration.

History of the identification of “Dark-belly”

The type species of the genus Tyrannochromis is T.macrostoma, first described by Regan (1922) in thegenus Haplochromis on the basis of one specimen of22.5 cm standard length (26 cm total length). Regan’sfigure is reproduced as fig. 43 in our book (Eccles &Trewavas, 1989). The Christy collection contained fiveother specimens of 12-20’cm total length. All thesehave white flanks below the mid-lateral black stripeand the markings above this stripe stand out on a palebackground. Dr. Digby Lewis and his collaborators(1986; p. 39) have a striking photograph of a fish la-belled Cyrtocara macrostoma with all the lower partof the flank black or dark brown, and their text re-marks on this peculiarity.

Could this be the same species as our white-belliedmuseum specimens? We had verbal reports of the abil-ity of this predator to change colour, but with no de-tails about the conditions in which this might occur.

Then, when our book was nearly finished, Ecclesfound in the Museum a specimen that had beenwrongly catalogued as H. polyodon. Its lower flankswere tinged brown. Here, he thought, is our “Dark-belly”, and as it was not either of the named speciesof Tyrannochromis he described it as a “new species”

with the name T. nigriventer, which means “black-belly”. The photograph of this fish is fig. 47 of ourbook (Eccles & Trewavas, 1989). The chief measur-able difference between this fish and the other speciesof Tyrannochromis is the short upper arm of the upperjaw (premaxillary pedicel). In T. nigriventer this isless than one third of the length of the head. In theother species it is very long, more than one third ofthe length of the head, enabling the fish to protrude itsmouth as a long funnel without dislocating its jaws.The photograph on p. 6 of Konings (1990a) shows themouth protruded, but not to its full extent. (This fishis a male in which the overall blue colour masks theadult pattern, of which the only indication consists ofa black mark on each scale, more prominent as theventral surface is approached, giving the flanks a soiledappearance).

The paragraphs on p. 103 of Eccles & Trewavas oncoloration and ecology apply to T. macrostoma andnot to T. nigriventer.

So were Dr. Digby Lewis and his collaborators mis-taken in identifying “Dark-belly” with T. macrostoma?This question was answered by our diving naturalists.

First, Andreas Spreinat of Göttingen caught and pre-served some “Dark-bellies” and also some other preda-tors. These he sent to the BMNH for our opinions.

38

A juvenile Tyrannochromis nigriventer photographed atChewere. The tips of the premaxillary pedicels bulge theskin clearly before the eyes. (Size approx. 10 cm TL)

Our book was then in the last stages, but after it waspublished Gordon Howes made some measurementsfor me. The pre-maxillary test showed that those la-belled “Dark-belly” by Spreinat agreed with T.macrostoma. The others were a good match for T.nigriventer. Spreinat also found that in his photographsthe ends of the premaxillary pedicels could be seenbulging the skin. His measurements of these, thoughnaturally inexact, gave a similar result, “Dark-belly”having pedicels whose length went 2.0 to 2.6 times inthe head length, whereas those of the other species,identified as N. fuscotaeniatus, went 3.5 to 3.75 times.The relationship between N. fuscotaeniatus and T.nigriventer will be mentioned below.

In November 1989 Ad Konings made a further visitto Lake Malawi and he too caught a number of “Dark-belly” and some of a species agreeing with T.nigriventer. The premaxillary test again pointed to T.macrostoma as the correct name for “Dark-belly”.

Both divers confirmed that the dark belly is the usuallive colour of adult, confident T. macrostoma, in thelake (except blue breeding males). Konings (1989; p.129) has a photograph of a guarding female that isvery dark except on the dorsal side, where the dorsaland dorso-lateral series of markings stand out clearlyon a pale background. This photograph is captionedT. maculiceps but Konings now believes, probably cor-rectly, that it is T. macrostoma. The adoption of blackcolouring when guarding young is reminiscent of thecolour changes in the mbuna Melanochromis auratusand may well be a parallel phenomenon. Koningsstates (1990a; p. 179) that the dark colour is not al-ways present in individuals transferred to a tank.Spreinat saw pale-bellied individuals that were “in-disposed” (letter of 24-6-90). It is understandable thata fish being caught for the pot or for preservation in amuseum is not merely “indisposed” but terrified.Hence our pale-bellied museum specimens.

The problem of the specific identify of “Dark-belly”is now therefore solved.

Konings touched on two other problems in the ge-nus Tyrannochromis. The first is the validity of thetwo other nominal species, T. polyodon (Trewavas,1935) and T. maculiceps (Ahl, 1927). Each of thesewas based on a single specimen. To synonymize T.polyodon it is necessary to show that the greater depthof its body comes within the range of variation of T.macrostoma. To synonymize T. maculiceps it must beshown that its narrow head and interorbital width, aswell as possibly its wider band of teeth at any givensize, all come within the range of a reasonable sam-ple of T. macrostoma from the southern part of the

lake. The type specimen of T. maculiceps was caughtat the extreme northeastern part of the lake, a smallfish of 163 mm standard length. Other specimens as-signed to this species are from Chilumba and NkhataBay. It may be that this is a species or subspecies com-mon in the north. Until these tests have been appliedthe validity of the names remains an open question.

The second problem is the relationship between T.nigriventer and Nimbochromis fuscotaeniatus. Theyresemble each other in proportions and dentition, butN. fuscotaeniatus has a pigment-pattern which is usu-ally much less regular than in the figured holotype(Regan, 1922 and Eccles & Trewavas, 1989; fig. 79),forming irregular blotches as in other species ofNimbochromis. No such tendency to form blotches isseen in the holotype of T. nigriventer, in which theventero-lateral brown patch is broader and continu-ous. Konings (1989; p. 222) remarks on the close re-semblance between these two species.

Konings does not like to include fuscotaeniatus in

A juvenile Tyrannochromis macrostoma photographed atChitande Island. This specimen is identified by the factthat the tips of the premaxillary pedicels bulge the skinbetween the eyes. (Size approx. 8 cm TL.)

39

A female Tyrannochromis nigriventer defending her offspring. The photo was taken at Chinyankwazi Island. It shows thata guarding female T. nigriventer can adopt a rather dark coloration on the belly. This female was collected for positiveidentification.

Nimbochromis because of an alleged difference inbreeding habits (Konings, 1989; p. 128). The eggs ofN. livingstonii are fertilized on the substrate, then takeninto the mouth (Konings, 1989; p. 219), whereas thoseof fuscotaeniatus are said to be fertilized in the mater-nal mouth. The reference for the latter statement is apaper by Dorenstouter (1982), but the identity of thefish in the photograph accompanying this paper isdoubtful and the statement requires confirmation.There are other differences between N. fuscotaeniatusand typical species of Nimbochromis and the inclu-sion of fuscotaeniatus requires further consideration.

References

DORENSTOUTER, C.F., (1982) Haplochromis fusco-taeniatus Regan, 1921. NVC Periodiek (DutchCichlid Ass.) April: 43.

ECCLES, D.H. & E. TREWAVAS, (1989) Malawian cichlidfishes. The classification of some haplochrominegenera. Lake Fish Movies. Herten, West Germany.335 pp.

KONINGS, A., (1989) Malawi cichlids in their naturalhabitat. Verduijn Cichlids, Netherlands. 303 pp.Illustr. in colour.

KONINGS, A., (1990a) Cichlids and all the other fishesof Lake Malawi. TFH Publications, Inc. NeptuneCity, New Jersey. 495 pp. Many illustr. in colour.

KONINGS, A. (1990b) The Malawi genus Tyran-nochromis. Cichlidae (Brit. Cichl. Ass.) Vol 11 (3),pp. 71-73.

LEWIS, D., P. REINTHAL, P. & J. TRENDALL (1986) A guideto the fishes of Lake Malawi national park. WorldWildlife Fund. Gland, Switzerland. 71 pp. illustr.in colour.

SPREINAT, A. (1990) Zur Revision der Haplochrominendes Malawisees nach Eccles & Trewavas. DATZ(Aquar. Terrar. Z.) (4); pp 245-248, illustr.

This article originally appeared in the January 1991 issueof Cichlidae, the periodical of the British CichlidAssociation. Details of the BCA may be obtained frompage 98.

40

Maravichromis (C.) liemi was previously placed inCaprichromis Eccles & Trewavas (1989). The reasonwas that it had a strongly inclined mouth, a broad lowerjaw and a specialized feeding behavior. Although it isfunctional to split large genera into a few smaller ones,monotypic or oligotypic genera should be avoidedwhen possible. The peculiar structure of the mouth ofliemi and orthognathus is related to the feedingbehavior but other species in the lake also have simi-lar mouth structures and comparable behavior. In thegenus Protomelas there are several, undescribed spe-cies (e.g P. sp. “Paedophage”) with a similar mouthstructure, but none of these or other species likeDiplotaxodon greenwoodi, have been put or will beput into a new genus or into Caprichromis. PersonallyI think the different mouth structure is insufficient justi-fication for erecting a new genus for these two species.

M. (C.) liemi is reported to steal eggs and larvae frommouthbrooding females (McKaye & Kocher, AnimalBehav. 31, 1983), specifically from P. pleurotaenia. Theauthors probably meant Nyassachromis microcephalusas P. pleurotaenia remains close to the bottom andmouthbrooding females are not found in schools.

I have observed M. (C.) liemi at many different lo-cations all round the lake and it seems likely that theirdiet is not restricted to larvae and eggs alone. I have

A male Maravichromis (Caprichromis) liemi in full breeding coloration photographed at Eccles Reef.

A subadult Maravichromis (Caprichromis) liemi.

Maravichromis (Caprichromis) liemi (McKaye & McKenzie, 1982)

observed several times that M. (C.) liemi attacked malecichlids as well, preferably those with parasites(Argulus) on the throat region. It is therefore possiblethat M. (C.) liemi feeds also, or even primarily, on theseparasitic invertebrates since fish with Argulus on theirthroat are very common.

M. (C.) liemi is normally found near rocks whereasM. (C.) orthognathus prefers the open sand. Breedingmales congregate into colonies and construct a sand-castle nest (more than 1 m in diameter) against a rock.Mouthbrooding females stay in schools and probablyrelease their fry in the rocky habitat.

Ad Konings

41

Sciaenochromis gracilis was described from threespecimens in the Christy collection. The total lengthof the largest specimen measured 217 mm. Some sin-gle specimens have been imported alive under tradenames such as “Torpedo” or “Dark Line Torpedo”.Recently, the export of S. gracilis has taken a morefrequent character. This is certainly caused by the ef-forts of Stuart Grant to comply with the increased de-mand.

The distribution of this rare species seems to be re-stricted to the southern part of the lake. Observationsabout their feeding and breeding behavior and abouttheir breeding coloration are not published yet.

The specific name (gracilis = slender) is aptly cho-sen. The narrowly pointed head with a slightly down-ward curved profile is a striking feature that remindsone of Nimbochromis linni. A band, consisting of smallspots, runs diagonally from the nape to the upper partof the tail (peduncle). In this respect it could be mis-taken for S. spilostichus or Maravichromis formosus.Maybe gracilis should be placed into Maravichromisas has been suggested before (Konings, 1989).

Its large mouth, its teeth and its shape presume apredatory lifestyle. The strong tail indicates a fastswimmer with ability to spurt.

A courting male Sciaenochromis gracilis photographed in the author’s aquarium.

Sciaenochromis gracilis (Trewavas, 1935)

Outside the breeding period S. gracilis is a livelyswimmer. Every now and then they suddenly stop andmove their eyes noticeably. Although S. gracilis be-haves rather peacefully we should not house them intoo small quarters. Their behavior changes during thebreeding period. Then S. gracilis is able to spurt overlarge distances without apparent effort.

Courting is rather impetuous and intensive, but thefemale is not hurt. Males defend a territory and con-struct (in my tank) a crater nest with a diameter ofabout 50 cm. The eggs are very small and are ferti-lized inside the female’s mouth. Since the eggs aresmall many fry can be expected. Females with a sizeof approximately 15 cm may release over 100 fry. Ju-veniles are, in comparison to other predators of simi-lar size, very small and slim. They grow rather slowly,probably because they show the same behavior as theadults, i.e. they frequently stop moving and search thearea with their eyes only. Food must then be broughtinto their mouths to let them eat! At a size of about 2-3 cm they defend small feeding territories and quarrelamong themselves.

S. gracilis grows to a maximum size of about 25’cmand is in shape, coloration, and behavior a valuableaddition to any large aquarium.

Peter Baasch

42

Otopharynx decorus is a small to medium sized cichlidwhich is found predominantly over the open sandy bot-toms of the lake. Its distribution encompasses probablythe entire lake because specimens have been collected atVua as well as in the southern part of the lake. The maxi-mum size of O. decorus is about 18 cm (total length);females are only a little bit smaller than males.

This sanddwelling species lives in small groups,usually numbering not more than 6 members. Solitaryindividuals are frequently observed, too. In Decem-ber, when I made the observations, there was some-times one male individual in the group that showedsome of its breeding coloration. Territorial males havenot been seen. Members of the genus Otopharynx usu-ally spawn near rocks. It is therefore possible that O.decorus also spawns close to a rocky coast and thatthe males which I was able to observe were foragingand not breeding.

The feeding behavior of O. decorus is rather char-acteristic of an insectivorous cichlid. With its large eyesit carefully screens the sand. Only after it has locatedsomething interesting does it bite into the substrate.Its food consists mainly of invertebrates which hide inthe sand. The teeth on the lower pharyngeal bone aresomewhat enlarged in the center, which is a common

A male Otopharynx decorus taken at a depth of approximately 18 meters in Senga Bay.

Otopharynx decorus (Trewavas, 1935)

feature in cichlids which live on crustaceans and in-sects. The feeding behavior closely resembles that ofMaravichromis sp. “Double Spot” (see next page).Both species have a small mouth with which they se-lectively pick their food from the substrate. Both spe-cies are found sympatrically.

The neutral coloration pattern consists of three ratherlarge blotches on the flank. These blotches are not al-ways distinct. When observed in the somewhat sub-dued light at a depth of 20 meters, the spots are almostinvisible. I have never seen them black as in O.(Ctenopharynx) nitidus with which it is commonlyfound sympatrically. The coloration of the body of O.decorus is like that of the sand – light beige.

Among the sanddwelling cichlids of the lake it israther easily recognized by the combination of thesmall mouth and the size of the fish. When observedin its natural environment, its behavior will immedi-ately allow an identification. Identification of aquariumor preserved specimens is simplified by the fact thatO. decorus has a high number of soft rays in the dorsalfin, 13 or 14, whereas most other Haplochromines haveless than 12. It has, furthermore, a high number ofscales on the flanks, higher than any other known Oto-pharynx.

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Maravichromis sp. “Double Spot” is a rather small,sand-dwelling cichlid. Its maximum size is known tobe less than 14 cm (total length). It has a rather widedistribution since it has been found on the east coastnear Fort Maguire, as well as near Mdoka in the north-ern part of the lake, and near Chintheche. It has, how-ever, never been seen in Senga Bay, although cichlidsare frequently collected there.

M. sp. “Double Spot” is characterized by a diago-nal row of three double-spots. In some specimens thespots are so large that they form a broad, almost solidblack band. This feature helped me decide to put it inthe genus Maravichromis and not in Otopharynx.

The “Double Spot” prefers the open sandy regionsand is usually found at between 5 and 30 meters depth.Most individuals forage on their own, rarely two orthree individuals are seen together. The coast near FortMaguire is inhabited by a rather dense population, al-though they never forage in groups or schools. At thetwo other locations where I have seen M. sp. “DoubleSpot”, only single individuals were found. The fish ofthe east coast population have a yellow coloration onthe body and especially on the fins, whereas speci-mens from the other locations are more silvery andhave colorless fins.

Maravichromis sp. “Double Spot” scrutinizes the sand for something edible (Fort Maguire).

Maravichromis sp. “Double Spot”

The mouth of M. sp. “Double Spot” is very smalland is utilized to pick small invertebrates from the sand.The substrate is therefore scrutinized for anything ed-ible. The characteristic posture of this cichlid is poisedover the sand, visually screening the area in front ofthe head with its large, movable eyes. The fish swimsfrom one place to the next and everywhere it halts andscrutinizes the sand. In this respect M. sp. “DoubleSpot” resembles Otopharynx decorus. The latter spe-cies also forages in small groups.

The specialized feeding behavior of M. sp. “Dou-ble Spot” might also be used for screening parasitesand fungus on the body and fins of sick fish. Once Iobserved a female Nyassachromis prostoma, with avisible fungus in the dorsal fin, who presented the ail-ing fin by laying itself on the sand in front of a forag-ing “Double Spot”. At the same moment a M.labidodon disturbed the scene, so I could not seewhether M. sp. “Double Spot” would actually cleanthe prostoma’s fin or not. It is, however, likely that itdoes, otherwise the ailing fish would not present itsfin in the way it did. The specific coloration pattern ofthe “Double Spot” is unique among the haplo-chromines of the lake and might function as a signalto other species that it is a “Cleaner”.

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Mpanga Rocks belong to a ridge which runs from themainland into the lake. It is about five kilometers off thecoast near Chilumba. At the beginning of the rainy sea-son, in December 1990, the water was very low and arather large portion of the upper boulders was above water.I had dived here in May 1989 and I wanted to observethe cichlid population at another time of the year. Afteran one-hour dive in the upper 20 meters of the rockyhabitat, which consists of huge boulders and large caves,I returned to the boat. Saulos Mwale had been diving aswell and told me that he had been to the sand and col-lected some new fish. I had never realized that the sandwas within reach and expected the rocks to plummet todepths beyond 60 meters. Saulos, without a depth gauge,estimated the depth of his new find at between 35 and 40meters. It is remarkable how he always knows the exactdepth, distance, and place without any technical device.When I dived for the sand, I found Nyassachromis sp.“Mpanga” at 36.4 meters! The sand at this site is rathercoarse and virtually void of the typical sand-dwellingcichlids you may find at other places. A regular and strongcurrent between the boulders of Mpanga Rocks may havetaken away all the fine material and left the coarser. Allspecies of Nyassachromis feed from the open water andnot from the sand. For them the structure of the substrate

A sand-turret nest of Nyassachromis sp. “Mpanga”.

A courting male Nyassachromis sp. “Mpanga”.

Nyassachromis sp. “Mpanga”

is therefore of less importance.Breeding males congregate into large colonies and

construct turret nests on the sandfloor. A photograph of atypical Nyassachromis nest is shown above. The dip ontop of the cone is the spawning-site. Such a dip is rarelyhorizontally placed; usually it tilts to one end. In this waythe females, which remain in schools in the open water,have a better view of the courting male when he displaysin the nest.

Mouthbrooding females stay in schools and may re-lease their broods simultaneously.

The size of adult N. sp. “Mpanga” ranges between7 and 10 cm.

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Otopharynx (C.) nitidus is a rather common sand-dwell-ing cichlid which is observed throughout the lake. Themaximum size measures around 15 cm for males; fe-males remain considerably smaller and were never seenlarger than 10 cm.

The preferred habitat is the open sand at depths below15 meters. Otopharynx (C.) nitidus is easily recognizedby the three large blotches, the yellow coloration on thelower part of the body, and the very large mouth. Themouth, though large, is weakly built and is meant to scooplarge amounts of the silty sediment which lies on the

A male Otopharynx (Ctenopharynx) nitidus in breeding coloration (Fort Maguire).

At some locations O. (C.) nitidus forages in large schools, sometimes numbering over a hundred individuals.

Otopharynx (Ctenopharynx) nitidus (Trewavas, 1935)

sand. The sediment is filtered for something edible,mainly invertebrates. In some areas large schools forageon the sand but mostly small groups of about five indi-viduals are more common. On only one occasion have Iseen males in breeding coloration (Fort Maguire; De-cember 1990). The few males I had observed seemed tohave a territory but a nest construction was not visible.Otopharynx (C.) intermedius, a closely related speciesbut without the yellow coloration, builds a sand-castlenest on a rock. The nitidus male follows females over arelatively large distance.

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When T. D. Iles described Copadichromis virginalis in1960 he pointed to the fact that natives distinguishedbetween the two morphs he regarded, at that time, asbelonging to this species. The natives called the twomorphs “Kaduna” and “Kajose”. The holotype of C.virginalis is of the Kaduna morph. The differences be-tween the two morphs are minimal. Kaduna has a smallersize, relatively larger eyes, and a deeper body and caudalpeduncle than Kajose and has little or no yellow pigmenton the body (females and non-breeding males) and usu-ally 16 spines in the dorsal fin. Kajose usually has 17dorsal spines and a yellowish coloration.

A male Copadichromis sp. “Virginalis Gome” at the entrance of its nest.

This is a characteristic nest of a Virginalis-type

Since its description C. virginalis has only sporadi-cally been observed in its natural environment, and thisapplies to both morphs. Now there is evidence that thetwo “morphs” represent two different species. In Decem-ber 1990 a population of both these species was found atthe same location but not in exactly the same habitat.The so-called “Fire-Crest Mloto” complies with the de-scription of the Kaduna and may therefore be conspecificwith C. virginalis. It is normally found, during the breed-ing period from August to November, at a depth of morethan 30 m. At the same location, Gome Rock (close tothe Mozambique border), a virtually identical species wasobserved. This species, for the time being termed C. sp.“Virginalis Gome” may be conspecific with the Kajose-Virginalis. It was found mainly at depths between 7 and15’meters.

C. sp. “Virginalis Gome” is larger, about 15 cm maxi-mum total length, than the “Fire-Crest Mloto”, whichhas a maximum size of about 12 cm. The females have alight, yellowish-brown coloration on the body and differalso in size from the smaller and silvery colored femalesof C. virginalis.

The only noticeable difference between the males ofboth species is the red colored band in the dorsal fin ofthe “Fire-Crest Mloto”. (cont. p. 47).

Copadichromis sp. “Virginalis Gome”Ad Konings

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At a depth of 30 meters the red color appears as ablack band in the dorsal of the “Fire-Crest” and femalesmay distinguish the two species by the pattern in this fin.Theoretically females of both species may live in mixedschools but this was not observed. Males of C. sp.“Virginalis Gome” were found to build their nests in agently sloping area of the coast whereas C. virginalismales constructed similar nests in the steeper part of thecoast and at significantly deeper levels. Such a nest con-sists of a crescent-shaped wall of sand which is posi-tioned under an overhanging rock or beneath a suspendedpart of a rock (see photo page 46). The nest is usually asbig as or bigger than the rock. It is important that therock forms the ceiling of the spawning-site. This type ofnest is constructed by all Copadichromis which have ablack male breeding coloration with a light colored flareon head and dorsal.

The assignment of the two species at Gome Rockto Kajose and Kaduna is relatively convincing; the situ-ation may differ at places where only one Virginalis-type inhabits the biotope or where only one type hasbeen found. This may be the case at Chitande Island.In November 1990 a population of a small Virginalis-like cichlid were inhabiting a wide depth-range of therocky coast. This species, C. sp. “Virginalis Chitande”,occurred at depths of between 5 and 30 meters. Males,

A male Copadichromis sp. “Virginalis Chitande” in full breeding coloration.

A female Copadichromis sp “Virginalis Chitande”.

with a maximum size of approximately 12 cm, have adark-blue breeding coloration with a light-blue flareon the upper part of the body. Females have a silverycoloration (see photo above) and were found at ratherdeep levels (about 25 m). The morphological featuresof “Virginalis Chitande” indicate that it is conspecificwith C. virginalis but the coloration (band in dorsal)and depth distribution does not agree with that of the“Fire-Crest Mloto”. Closer examination is needed toassign “Virginalis Chitande” to one of the known spe-cies or to describe it as a new one.

Copadichromis sp. “Virginalis Chitande”Ad Konings

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This unusual cichlid was found on the east coast ofLake Malawi, just south of the Mozambique bor-der, at such places as Nurungu and Fort Maguire.At first glance this cichlid appears to belong to thegenus Diplotaxodon. For when first observed it wasseen lying in a large pile of freshly caughtDiplotaxodon species. After analyzing this cichlidin a more detailed fashion, it seems probable that itdoes not belong to the genus Diplotaxodon. Its ap-pearance also reminds one of a Rhamphochromisspecies, although it is doubtful that it belongs tothat genus either. Of the two genera, it definitely ismore closely related to the genus Diplotaxodon.Since there are not any known cichlid genera inLake Malawi that appear to be half-way betweenRhamphochromis and Diplotaxodon, it seems prob-able that this cichlid represents a new genus ofMalawi cichlid. Only more detailed scientific analy-sis will be able to determine to which it rightfullybelongs. The head and mouth of this fish are enor-mous in comparison with the rest of its body. This,along with its large, highly visible teeth, would seemto indicate that this cichlid is piscivorous. The un-paired fins are reduced in size (except for the anal,which is unusually large for a piscivore), similar to

The author collected “Diplotaxodon” sp. “Big Head” near Fort Maguire. Photo Ad Konings.

“Diplotaxodon” sp. “Big Head”

that found in Rhamphochromis, presumably indi-cating that this fish is a fast swimmer.

“Diplotaxodon” sp. “Big Head” was caught onhook and line using the lake sardine, Engraulicyprissardella, as bait. They were caught several hundredmeters offshore along with species of Diplotaxodon,Alticorpus and Rhamphochromis. Its large eyes andlack of significant coloration indicate that this fishis most likely to frequent deep water. The largestspecimen observed was about 35 cm in total length.Very little else is known about this interesting look-ing discovery, and, like virtually all the cichlids inLake Malawi, it undoubtedly is a mouthbrooder.

Mark Smith

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This fascinating Lethrinops species appears to beundescribed and I, therefore, gave it the non-scien-tific name “Christyi Fort Maguire”. I found thespecimen in the photograph at the bottom of a largestack of nets in a native fisherman’s dug-out canoe.The locality was Fort Maguire, just south of theMozambique border.

A method of fishing that some Malawian fisher-men utilize is the draping of a large net, a so-called“Chirimila”, down into the open waters between twocanoes. The larger nets extend down to beyond 70meters (220 feet). The nets in the canoe in which Ifound the new Lethrinops were quite lengthy andthe cichlids caught sported a characteristic deepdwelling cichlid coloration of black vertical bar-ring on a silvery gray body.

Lethrinops sp. “Christyi Fort Maguire” wascaught along with L. gossei in the same net. Theextreme slope of the upper head profile and pointedsnout would seem to indicate that this deep dwell-ing Lethrinops pokes its snout into the muddy bot-toms of its biotope in search of food.

Because of the metallic green coloration on thehead and snout, it must be a male. It superficiallyresembles L. christyi; however, L. christyi has a

Lethrinops sp. “Christyi Fort Maguire” seems to be a very interesting species from the deep waters.

Lethrinops sp. “Christyi Fort Maguire”

more elongated snout than does this undescribedLethrinops. It also seems closely related to L.macracanthus, although the snout of the latterdoes not protrude outward as much as does that ofL. sp. “Christyi Fort Maguire”. Other Lethrinops,such as L. altus and L. mylodon, also appear to beclosely related to this undescribed species. Thesnout of L. altus is more elongate than that of thisspecies while L. mylodon’s body is stouter.

Only one specimen, with an approximate lengthof 15 cm, was found in the fishermen’s nets, so it ispossible that the fish is either rather rare or is notnormally found in the area and or depth at which itwas caught. It was in any case remarkable that aspecies of Lethrinops, usually a bottom-dwellingcichlid, got into a net which is specifically used tocatch “Utaka” and other open water cichlids. Thenet may have been dragged over the bottom becauseit is unlikely that a Lethrinops with such a mouth-structure lives in the open water. The length of thenet and the lines was difficult to estimate but wascertainly more than 70 meters.

Mark Smith

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The genus Taeniolethrinops is characterized by spe-cies which have a diagonal band on the body and havea Lethrinops-type mouth. If we regard T. laticeps assynonymous T. praeorbitalis, three species are knownto belong to this genus. The Lethrinops-like cichlidsare very difficult to distinguish from each other as theyall have a similar body-shape and are silvery or sand-colored when not breeding. The best way to tell thesecichlids apart, without dissecting specimens, is to waittill they are in their breeding period. Males usually ac-quire the most fantastic breeding colorations and can thenbe identified. The type of habitat and type of nest givefurther indications of identity. Before we know whatbreeding coloration belongs to which species we have toobserve, photograph, and collect each species. A closeexamination of a few preserved specimens then tells uswhether we are dealing with a scientifically known orwith a new species. This procedure takes, of course, a lotof time because breeding in most Lethrinops and alliedspecies occurs only at certain periods of the year.

In November 1990 we found several aquaristicallyunknown species clearly belonging to the Lethrinopsgroup of sand-dwelling cichlids. The species which isshown in the picture above was found in very smallnumbers near a village called Ntekete on the east coast.

A breeding male Taeniolethrinops sp. “Furcicauda Ntekete” patrolling its territory over the sand.

A female Taeniolethrinops sp. “Furcicauda Ntekete”(Ntekete, east coast).

Taeniolethrinops sp. “Furcicauda Ntekete”

A breeding male of T. sp. “Furcicauda Ntekete” con-structs a large craternest (approx. 100 cm diameter)in the sand with an elevated rim around it. Two nestswere seen built at a depth of about 15 meters. Fe-males were found solitary, foraging on the sand inthe vicinity of the male’s nest.

The two specimens Saulos caught for me proved tobelong to a new species. It has, however, some simi-larity to T. furcicauda, from which it differs by havinga shorter snout and a larger eye. T. sp. “FurcicaudaNtekete” grows to a size of about 20 cm.

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Lethrinops sp. “Longipinnis Ntekete” was only foundin small numbers in shallow water on the east coastnear the village of Ntekete. Breeding males were ob-served defending their peculiar nests in November1990. Males, which have a maximum size of approxi-mately 19 cm, construct a type of sand-castle nestwhich is unique among Malawian haplochromines. Thenest consists of a large crater dug in the sandy bottomand an elevated wall made of five to eight large tur-rets. These turrets are about 30 cm high and about50’cm wide at the base. The diameter of the completenest is about two meters! Several of these nests wereobserved in 2 to 3 m of water. Females were solitaryand foraged on the sand.

The only other Malawian species which builds turretsaround the spawning-site is L. auritus (Lewis, 1980: J.Sci. Tech. Malawi; 1 (1); pp. 36-37). L. auritus, the small-est member of the genus, builds its nest in muddy envi-ronments and does not dig a crater but heaps small tur-rets in a circle around a centrally located heap, alongsidewhich it digs a shallow pit for a spawning-site.

L. sp. “Longipinnis Ntekete” resembles L. longipinnisin appearance but has a longer snout, a shallower body,and shorter pectoral fins than this species, which is knownonly from the southern part of the lake.

A male Lethrinops sp. “Longipinnis Ntekete” at its nest.

Above: The size of the nest with the turrets is enormous.Below: L. sp. “Longipinnis Ntekete” has a magnificentbreeding coloration.

Lethrinops sp. “Longipinnis Ntekete”Ad Konings

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The genus Gephyrochromis contains two scientificallydescribed species and two undescribed ones. The de-scribed species are G. moorii and G. lawsii. G. mooriiwas exported from the lake in the late seventies and al-though it has the most beautiful coloration among thefour species – especially the bright orange color on thethroat – it never became a very popular cichlid. The rea-son may be that it is rather difficult to collect as it lives atdeep levels. For reasons unknown to the author it is called“Sergeant” by the local fishermen. It is not known wherethe fish were collected but G. moorii has been observednear Msuli Point (Chintheche) at a depth of about 40meters. Like the other species of the genus it lives oversand, preferably in areas which are covered with a layerof silt or mud.

G. lawsii has been observed in the bay near Ruarwe ata depth of about 25 m. They live in small groups andforage over the muddy floor of the bay. They have re-cently been exported from the Mbamba Bay, Tanzania.Another species of the genus is also found in the MbambaBay; it has been named G. sp. “Liuli” for the time being.It lives in rather shallow water and is sometimes caughtin the nets of the fishermen on the beach (Peter Knabe,pers. comm.). This species has also been exported re-cently by Hans Fleischer, who collected it near Liuli.

The fourth species in the genus, G. sp. “Zebroides”, is

Gephyrochromis sp. “Zebroides” has a very attractive coloration pattern.

A male Gephyrochromis sp. “Liuli”.

Gephyrochromis

found on the east coast near Masinje and Fort Maguirebut is also observed in the bay near Ruarwe. It has aremarkable color pattern, unlike that of the other threespecies. G. sp. “Zebroides” was observed at depths rang-ing from 7 to 30 meters. They occur in small groups orare solitary. Their foraging grounds are on the sand butbreeding may take place in the rocky habitat. Some speci-mens were observed among the rocks and had a verydark coloration which almost obscured the regular pat-tern of bars. A distinction between males and females isnot possible on the basis of coloration alone.

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The species in the Pseudotropheus zebra complex forma closely related group of rock-dwelling Mbuna. Theyprovide the ichthyologists with many questions andmore problems. P. zebra, of which the exact type-lo-cality is unknown, is a polymorphic and variable spe-cies. In several populations there are so-called OB-morphs (orange-blotched, i.e. orange body color withblack blotches of variable sizes) and O-morphs (withan entirely orange-colored body). Some populationsdiffer from the holotype, which has distinct, dark ver-tical bars, by having no vertical bars at all. But such apopulation can also have OB- and O-morphs (e.g. thepopulation at Jalo Reef; pers. obs.). Furthermore thereare populations in which the males have a bright reddorsal fin. It is still questionable whether these red-dorsal populations belong to P. zebra or form anotherspecies because OB-morphs have never been foundamong them. Another species, P. sp. “Zebra Cobalt”,lacks vertical bars on the body, lives sympatrically withP. zebra and also shows OB-polymorphism. How canone distinguish between a non-barred P. zebra popu-lation and the “Zebra Cobalt”?

The region with the most complicated communityof P. zebra related species is that around the MaleriIslands in the southern part of the lake. Here, five dis-

Pseudotropheus sp. “Zebra Charo” is a very common Mbuna at Charo.

Pseudotropheus sp. “Zebra Charo”

tinct species inhabit the different rocky habitats. Oneof the species, P. sp. “Zebra Black Dorsal”, is easilyrecognized by the black band in the dorsal fin. It re-sembles P. heteropictus from Thumbi West Island andmight even be conspecific with this species.

At Charo and Kakusa we find no less than four spe-cies of the P. zebra complex. These two rocky coasts,which are about 7 km apart, are the only known habi-tats of P. sp. “Zebra Charo”. This Zebra is easily rec-ognized by the black band in the dorsal fin which couldindicate some relationship with P. heteropictus fromthe south. The latter species, however, lives in the sedi-ment-rich rocky biotopes as does P. sp. “Zebra BlackDorsal”. P. sp. “Zebra Charo” lives in the sediment-free rocky regions and behaves more like a member ofthe genus Cynotilapia. It does not have unicuspid teeth,a prerequisite to belonging to that genus. Males arecharacterized by an orange-colored trailing edge in thedorsal fin. The population at Kakusa has less colorfulmales but clearly belongs to the same species.

P. sp. “Zebra Charo” shares the habitat with P. sp.“Zebra Cobalt” (Pearl variety), P. sp. “ZebraGoldbreast”, and to a lesser extent with P. sp. “ZebraGold”, as this species is normally found at deeper lev-els of the biotope.

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Recently, there have been several new cichlid im-ports from Lake Victoria. One of the most colorfulimports is a haplochromine species, nicknamed“Flameback”. Flameback haplochromines are foundin shallow, near shore waters along the northerncoast of Lake Victoria and have been reported alongthe south as well.

Males in breeding dress exhibit scarlet red fromthe forehead dorsally to the caudal peduncle, witha turquoise-blue coloration from the lateral line

“Haplochromis” sp. “Flameback” is one of the most colorful cichlids of Lake Victoria.

VICTORIANCICHLIDS

Exciting new discoveries

Laif DeMason

ventrally along the body to the belly region. Thereis faint darker blue barring along this body area.Theanterior part of the dorsal fin is light gray becom-ing scarlet posteriorly along the soft rays. The cau-dal fin is scarlet, the anal fin orange, and both pel-vic fins are jet black.

Females are mostly silvery gray-green with a faintpink sheen dorsally and yellow anal fins. The fe-males are mouthbrooders with approximately 25offspring per brood.

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“Haplochromis” sp. “Migori” is another very interesting find. Photos by Ad Konings.

Unfortunately, not much is known about the hab-its of this wonderful fish. It appears to eat plank-tonic particles from the water column and will eatany aquarium fare.

Other interesting haplochromines are also beingcollected and sold from waterways near Lake Vic-toria, but not actually from the lake proper. Sincesome of these species are exported by commercialfirms, exact collecting localities are not alwaysknown. Such is the case with an interestinghaplochromine species named “Migori”. “Haplo-chromis” sp. “Migori” is reportedly from the MigoriRiver, which feeds Lake Victoria from the east. In-

formation is not available on the exact habitats ofcommercial collections.

“Migori” males have a gold-green body with redpatches on the operculum, and behind the pectoralfins. The dorsal and anal fins are suffused with alight blue hue, which is sometimes visible on thebody adjacent to the anal and caudal fin. There aretwo to five yellow-orange spots present in the analfin as well. The head is grayish with a slightly con-vex shaped forehead. Usually prominent is a black

bar between the eye and corner of the mouth, typi-cal of many Victorian haplochromines.

The female “Migori” is yellow or yellow-greenin coloration, with a thin black longitudinal stripefrom the operculum to the caudal. The ventral sur-face, from the mouth to the tail, is clear yellow incolor as are the ventral fins and the soft rayed por-tion of the dorsal fin. Females are mouthbrooders,having more than twenty offspring per brood.

A haplochromine with similar male colorationfrom the northern shores of Lake Victoria is alsoknown. It is unclear whether these two cichlids arethe same species. Until systematics and distribu-

tions for haplochromine species new to theaquarium hobby are known, an exact classificationis impossible.

A third colorful import is the true Astatotilapianubila. In the past many fishes sold as Haplochromisnubilus were only similar and varied haplo-chromines from Lake Victoria. The true nubila maleis a velvety black fish with a bright red edge to thedorsal fin, along with a bright red caudal fin and abright red anal fin. The latter is decorated with sev-

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“Haplochromis” sp. “Flameback”, a female.

A male Astatotilapia nubila.

“Haplochromis” sp. “Migori”, a female.

eral golden-yellow spots. A. nubila is another shal-low-water species found over sandy bottoms at sev-eral places in the lake. The females are mouth-brooders and have between 25 and 50 progeny ineach brood.

With these new haplochromine imports fromLake Victoria, interest in these fascinating cichlidscan be revitalized, making more commercial col-lection a viable pursuit. Like Lake Malawi, LakeVictoria has provided a site for complex speciesflock evolution. Both trophic and habitat spe-cializations evolved gradually to give us the present-day lineages of species.

Anatomical features, such as dentition, shape ofjaw, and morphology of skull parts, have been re-fined in each species (Greenwood, 1981).

Male coloration is probably a major factor in theprevention of hybridization. In many cases, speciesare virtually identical in their anatomy and can bedistinguished only by male breeding colors. Thisslows commercial collection considerably and in-creases the cost the importer must bear.

With breeding colonies of colorful cichlids suchas “Flameback” and “Migori” in place, progeny are

now available. With increased interest in Lake Vic-toria fishes, many additional species will becomeavailable.

We have only scratched the surface of the manywonderful and colorful varieties from Lake Victo-ria. Hopefully many new and exciting Victoriancichlids will soon be available.

References

FRYER, G. & T.D. ILES (1972) The cichlid fishes of thegreat lakes of Africa. Oliver & Boyd, London &Edinburgh.

GREENWOOD, P. H. (1981) The haplochromine fishes ofthe east African lakes. Kraus International Publica-tions, München.

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The species in the genus Chromidotilapia, which aremouthbrooders, belong to a still relatively unknowngroup of cichlids and are rarely kept in the aquariumsof the West African “cichlidists”. This is remarkablesince this genus offers several species which have de-veloped various and highly advanced breeding strate-gies. Moreover there are many Chromidotilapia withan attractive coloration and which are easy to keep ina community aquarium with other West Africancichlids such as those from the genera Pelvicachromisand Parananochromis.

A female Chromidotilapia sp. “Mondemba”. Photo by Roland Numrich.

WEST AFRICANCICHLIDS

Two new Chromidotilapia species

Roland Numrich

The species in Chromidotilapia can be split into twodifferent groups:

(1) The guntheri – kingsleyae group with the spe-cies C. guntheri and subspecies C. guntheri loenbergiand C. guntheri bosumtwensis and C. kingsleyae plusabout six undescribed species from the Ogoowe drain-age in Gabun and two species from Congo.

(2) The batesii – finleyi group with the species C.finleyi, C. batesii, C. linkei and an undescribed spe-cies from the Korup region.

The features of the finleyi-batesii group make these

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A male Chromidotilapia sp. “Mondemba”. Photos by Roland Numrich.

fishes excellent aquarium inhabitants. They are rela-tively small – a maximum size of approximately 10 to12 cm – and there are a number of attractively coloredraces known, e.g. the “Mungo” variant of C. finleyi orthe “Loum” variant of C. linkei.

The smaller species of the guntheri-kingsleyaegroup, in contrast to the larger species of the samegroup, exist in many different color variants in the re-spective drainages of their distribution. A similar situ-ation is also found in the usually sympatric genusPelvicachromis.

Here I introduce a newly imported species of thethe batesii-finleyi group. Chromidotilapia sp.“Mondemba”, as it is called, is found in the river sys-tem at the border between Nigeria and Cameroon andis new to science and to the hobby. C. sp. “Mondemba”grows to a length of approximately 7 cm and resem-bles C. batesii, although it has also some features incommon with C. finleyi. Due to the lack of ample pre-served or live material its status as a valid species isstill unresolved.

The borderland between Nigeria and Cameroon was,a couple of years ago, completely unknown territorysince it consists of virtually inaccessible mountainousterrain. The natural borderline is formed by the Akpa-

Yafe River which is blessed with a rich and varied floraand fauna, of which many species are endemic to thisregion. The hinterland of the approximately 160 kmlong river forms one of the last intact rainforests inWest Africa, which is in the process of being protectedas a national park by both countries. In the processseveral surveys have been made and it was estimatedthat more than 140 species of fish live endemic in thisregion!

The first specimen of C. sp. “Mondemba” was col-lected in a small stream in the eastern Oban Hills in1980. In 1987 more specimens were found in the Akpa-Yafe rivermouth near Ikono. In 1989 the first speci-mens were collected and brought back alive to Ger-many by M. Freier from Würzburg. He caught the fishat the western slopes of the Rumpi Hills near theCameroon city Mondemba (Ndian drainage).

The fact that from the rivers east of the Rumpi Hillsseveral populations of C. finleyi are known and in Ni-geria west of the Oban Hills only C. guntheri, mayindicate that this new species is endemic to the region.However, it is possible that C. sp. “Mondemba” alsolives in the upper Munaya River (Cross River system).

Keeping C. sp. “Mondemba” proved to be rathersimple as they are not aggressive towards each other

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The collection site of Chromidotilapia sp. “Shiloango”

and eat any type of aquarium fare. This situationchanged when the fish reached sexual maturity. I hadput a male and female in a separate tank in order tobreed them. The male became too aggressive and Ihad to separate the pair with a divider until the femalewas ready to spawn. Although the pair spawned sev-eral times they did not succeed in raising the offspringbecause the female repeatedly swallowed the eggsduring the night after the spawning. I kept the pair inwater with a pH of 5.5 and a conductivity of 120microSiemens. All we know of their breeding tech-nique is that C. sp. “Mondemba” is a ovophilousmouthbrooder. The reason that the brood was swal-

lowed could be that the handing(mouthing) over of the eggs to themale did not function properly as isknown for several aquarium po-pulations of C. finleyi as well.

In contrast to the small species ofthe batesii-finleyi group, the Chro-midotilapias from the African stateGabun, with its main river Ogoowe,are much larger. Fully mature fe-males of C. kingsleyae can reach asize of about 20 cm. For most of thescientifically undescribed but aqua-ristically known species from thisarea the maximum size lies around15 cm. A main feature of these spe-cies is the pointed, somewhat down-wards directed mouth, which indi-cates a feeding behavior similar tothat known for species of the SouthAmerican genera Geophagus andSatanoperca. The members of theguntheri-kingsleyae group are sub-strate-sifters, i.e. they forage by“chewing” the substrate and passingit through the gills retaining any ed-ible matter.

In the summer of 1988, during anexpedition in Gabun, the author andF. Bitter from Lünen were able tocollect five undescribed species andC. kingsleyae.

The second species I would liketo introduce here was collected in thecoastal region in the southwesternpart of Gabun close to the border ofCongo. It was found in the Nyangadrainage. I have chosen to name it

Chromidotilapia sp. “Shiloango” as it may beconspecific with the species from the Shiloango thatLinke & Staeck (1980) mentioned. These authors prob-ably derived their data regarding the distribution ofthis mouthbrooder from the ample material in theMusée Royal de l’Afrique Central in Tervuren, Bel-gium. Their collection consists mainly of fish collectedin the Niari-Kiolu and the Loango River systems inthe, nowadays, Republic of Congo. Some specimens,however, were caught in the Nyanga in Gabun andappear, in spite of the wide distribution this assumes,conspecific with the Congo populations.

We found C. sp. “Shiloango” in some tributaries of

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Chromidotilapia sp. “Shiloango”. Photo by Roland

The distribution of Chromidotilapia sp. “Mondemba” (blue) and C. sp. “Shiloango” (red).

the Nyanga River in the savannas as well as in thecoastal rainforests where we caught them in a numberof different streams. In the main drainage system inthis region, the Ngounie River between Du ChailluMassiv and the coast, we find only C. kingsleyae.

The habitat of C. sp. “Shiloango”, which grows to amaximum size of about 12 cm, consists of small riv-ers and fast flowing streams with bottoms of light andfine sand. Aquatic plants are seldom seen but here andthere a waterlily (Crinum spec.), with two-meter-long,floating leaves, gives protection to the many killifishesof the genus Plataplochilus. The Chromidotilapiasforage in loose groups of up to 12 individuals in theshallow water close to inundated banks, and dive forshelter in the leaves and root-systems which form athick carpet in theseareas. It was thereforedifficult to catch spe-cimens and we finallyrelied on the localfishermen whocaught some withhook and line (thismethod proved veryeffective when wecollected some small-er Parananochromisin the northern part ofGabun).

The composition of the water is, despite its proxim-ity to the coast, very soft and acid. We measured a pHbetween 5.2 and 5.8 and a conductivity of 20 to 35microSiemens. The temperature was rather low at 22°C. The main food of C. sp. “Shiloango” probably con-sists of the many 5 to 10 mm long, black-and-redcolored shrimps which resemble to some extent theAsiatic Zebrashrimps of the genus Caridina, but have,instead of bands, black blotches.

Breeding this Chromidotilapia has been successful,even in hard water. The “Shiloango” Chromidotilapiais an ovophilous mouthbrooder in which the male –assiduously – takes care of the brood. He defends thefry against the other fish in the aquarium till they havereached a size of about one centimeter. The youngChromidotilapias reach sexual maturity and start pair-ing-off at a length of about 4 cm!

References

MARTIN, C. (1989) Die Regenwälder Westafrikas.Basel.

REID, G. M. (1989) The living waters of southern Koruprainforest. WWF Report, number 3206/A8:1.

STAECK, W. & H. LINKE (1980) Afrikanische CichlidenI. Buntbarsche aus Westafrika. Tetra Verlag, Melle.

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All known Steatocranus species are found in therapids of the lower Congo or Zaïre River and itstributaries. Apparently Steatocranus ubanguiensiswas first collected during a scientific expedition tothe Mbomou River, a tributary of the Ubangui River,near Gozobangui. However S. ubanguiensis appearsto be found in the Congo (Zaïre) River as well, forit has also been found sympatrically with S.casuarius in the Malebo Pool at Kinshasa, the lo-cation where most Congo (Zaïre) River fish arecollected and exported. In careful looking throughthe hundreds of “Buffalo Head Cichlids” (S.casuarius) that make their way each month intowholesaler aquariums, one is guaranteed to findmany “contaminants” mixed in with S. casuarius,and S. ubanguiensis is often one of them.

In the wild, S. ubanguiensis does not seem togrow much larger than 6 to 7 cm in total length, butcan probably attain a larger size in captivity. ThisSteatocranus also behaves in a more aggressivefashion than does S. casuarius. The small to non-existent swimbladder and short stubby ventral fins,reminiscent of the suction disk-like ventral fins seenon some goby species, indicate that this cichlid in-habits fast flowing water. Even its squared-off,bluntly shaped head shows that it is used as an aidto cut through the currents of swift flowing water.

S. ubanguiensis’ chisel-like teeth are used to pickaway at the algae and micro-organisms encrustedon the rocks of its habitat. Yet, as with most othercichlids, S. ubanguiensis readily adapts to estab-lished aquarium fish food with great enthusiasm.

Steatocranus ubanguiensis. Steatocranus mpozoensis. Photos by Mark Smith.

Steatocranus ubanguiensis

Steatocranus mpozoensis is another species that isregularly found in shipments of S. casuarius out ofZaïre, West Africa. At first glance one may be de-ceived into assuming that S. mpozoensis is in facta juvenile S. casuarius. But upon closer examina-tion a couple of obvious differences stand out. Firstof all, the head of S. mpozoensis slopes more gradu-ally than does the blunt, blockhead-like appearanceof S. casuarius. Secondly, the fright pattern is verydifferent in S. mpozoensis to that in S. casuarius. S.mpozoensis has two horizontal, dark colored linesrunning down the length of the body, overlaying adark, checker-like pattern. Only the dark checkeredpattern is seen in S. casuarius. These dark mark-ings vary from one species to the next within thegenus Steatocranus and recognizing these varyingpatterns may help to identify the species.

This particular Steatocranus seems to behave ina manner similar to S. casuarius with little varia-tion.

S. mpozoensis was originally collected in theMpozo River, a tributary of the Lower Congo(Zaïre) River. What is remarkable about the MpozoRiver is that the pH of the water measured 8.5 to9.0 at the time of collection on September 1st, 1973.

Reference

ROBERTS, T. R. & D.J. STEWART (1976) An ecologi-cal and systematic survey of fishes in the rapidsof the lower Zaire or Congo River. Bull. Mus.Comp. Zool. 147, pp 239-317.

Steatocranus mpozoensisMark Smith

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Pelvicachromis taeniatus (Boulenger, 1901)

Pelvicachromis taeniatus, a male from Kumba Funge, Cameroon. Photo by Roland Numrich.

Pelvicachromis taeniatus, a female from Kumba Funge.

Pelvicachromis taeniatus from Calabar. Photo by Numrich.

P. taeniatus, a female from the Nyong River population.

P. taeniatus, a female from Calabar. Photo by Numrich.

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Pelvicachromis taeniatus (Boulenger, 1901)

Pelvicachromis taeniatus, the so-called “Red Taeniatus”. A male photographed by Roland Numrich.

Pelvicachromis taeniatus, a female of the “Red” variety. Photo by Roland Numrich.

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Diving among the boulders, I suddenly observedgreen dashes crossing below me. I watched themamong the rocks swimming at great speed againstthe strong current. I was surprised how fast a cichlidcould swim that way without an apparent, big ef-fort. From time to time an individual stoppedabrubtly and remained still for a few seconds,looked around, then continued racing. The person-ality of the elongated cichlid I was watching wasfantastic, and they didn’t look like they were rushingbecause of my presence, which they seemed to ignore.

Paraneetroplus nebuliferum in breeding coloration. Photo by Juan Miguel Artigas

CENTRAL AMERICANCICHLIDS

The re-discovery of Paraneetroplus nebuliferum

Juan Miguel Artigas Azas

This event took place in April 1990, just outsidethe town of Santo Domingo de Ocampo (95° 4’ W.Lon., 17° 09’ N. Lat.) in the northern part of theisthmus of Tehuantepec in Mexico. I was at Rio DosCaños, a small tributary of the Rio San Juan, whichbelongs to the Papaloapan river system in the Mexi-can state of Veracruz. I was about 45 km south fromCatemaco crater lake, well known as the place ofthe witches.

The water, with a pH of 8.0, GH of 4° and a KHof 5°, was yellowish and warm (over 28° C). The

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P. nebuliferum in normal dress. Photos by J. M. Artigas A pair P. nebuliferum.

river, at the site, was an average of ten meterswidewith perhaps ten meters of exposed riverbeds be-tween the banks. Because of the fairly steep gradi-ent, the water flowed at a moderate speed. Boul-ders were present on the bottom of the river whichwas about one meter deep. At some deeper spots itwas two meters deep. Pools had muddy to sandybottoms.

The cichlid I observed was indeed a cichlid ofthe genus Paraneetroplus. It was the Rio Papa-luapan equivalent of Paraneetroplus bulleri Regan,1905 of Rio Coatzacoalcos. I thought, it had to bethe long searched for P. nebuliferum (Günther,1860), a fish everybody interested in Central Ameri-cans has been asking me for, whenever I had col-lected in the Isthmus.

The type locality, as stated in the description, isan eastern lowland stream of the Isthmus ofTehuantepec, more precisely, at San Juan Evan-gelista. The last name makes it easy to find the ex-act spot; it is a small town located in the state ofVeracruz at the edge of the Rio San Juan. I wassnorkeling in the same river, not far away from thistown.

I spoke with a fisherman around Rio Dos Cañosand he told me that the fish is commonly known as“Corrientera”, referring to its habit of living in cur-rents. The same name is used for P. bulleri aroundRio Coatzacoalcos. The fisherman also told me thatthe cichlid was hard to get, but, despite its littlemeat, it was tasty and very much appreciated.

The accompanying pictures give some indicationof the coloration. The main characteristic is a darkhorizontal stripe over the entire body. The tail fin islarge and powerful and is an adaptation for swim-ming in currents.

Although its original description states a lengthof 35 cm, the larger specimens I saw could have

been around 25 cm. When the fish grows larger, thegreen color on the scales of the flanks intensify inbrightness. Large males show more spots which arealso present on the forehead.

In Rio Dos Caños I found P. nebuliferum in thecompany of three other cichlid species. One was aspecies of the Thorichthys group, called “Chonga”by local fishermen. It was abundant near the banksand in calmer water. It occurred mainly over muddyand sandy bottoms. In small numbers, “Cichla-soma” (Parapetenia) salvini with a beautiful col-oration was most frequently seen in quieter regionsof the river. Its local name is “Mojarra Pico deGallo” (cock’s comb cichlid). The deeper areas ofthe river were dominated by “C.” (Theraps)fenestratum, called “Testa Colorada” (red head) bylocals.

P. nebuliferum feeds on the lush algae whichcover the boulders in the stream. Its rather power-ful mouth faces downwards and tears off pieces ofthe algae. In this respect it has no competition fromother cichlids. Only “C.” fenestratum occasion-ally feeds on algae but it prefers to do so in thecalmer waters near the bank. It also has a moreomnivorous diet.

At the time of my visit, the end of the dry sea-son, all cichlid species were found in breeding ac-tivity. P. nebuliferum pairs were seen in the rapids,mainly in the shade of large trees. Females roundedup their fry, numbering up to two hundred, in closecircles, meanwhile grazing from the algae on therocks. Males rounded up the fry too, but more froma distance. They kept intruders away from the fry.Once in a while the pair would remain still in thestrong current while hovering over their fry. I no-ticed that at the slightest sign of danger, the frywould immediately scatter and get protection un-der the rocks. The female then stays at a safe dis-

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A Rio San Juan affluent, the collection site of Paraneetroplus nebuliferum. Photo by Juan Miguel Artigas Azas.

tance. Once danger had passed, the fry come outfrom their shelter and get all together again follow-ing the signals of the mother, who vigorously shakesher fins to her offspring.

Collecting some individuals proved quite a task,as this fish is very fast and intelligent. During aholiday afternoon, when people of the town weretaking a picnic at the riverside and kids had beenswimming all day long in the muddy pools, thewater was very murky and the fish did not see thenet. So, a friend of mine and myself, using castingnets, were able to collect three adult specimens, butonly after hundreds of throws. We had to disentan-gle the net from the rocks every other throw, withthe net getting torn in the process. After a while itbecame dark and we had to return the next day. Thewater was clear this time. The visibility was aboutthree meters and the “Corrienteras” very hard tocatch with the casting nets. I decided to snorkel forfry with a handnet. This proved to be a good choice,despite the fact that small Paraneetroplus nebuli-ferum are difficult to get this way because of theiraforementioned habit of scattering for protectionbeneath the rocks. It was hard to handle the handnetin the current, but I was able to catch some forty

small individuals.For taking adult fish home we used large plastic

jugs of fifty liters capacity. They were about halffilled. We took care that the water did not get toowarm, which is the worst type of danger since itwill deplete the oxygen and kill the fish. The frywere transported in a bucket closed with a lid. Theywere fed algae. Twice a day the water was changed.An extra step we usually take is to add a commer-cial tranquilizer to the containers of the large fish.This prevents stress and oxygen consuming fightsamong them. Given such treatment, cichlids havesurvived eight-days-long trips. This time all the“Corrienteras” arrived home safely.

Once home I placed the nebuliferum in quaran-tine using a commercial parasite killing solution inorder to prevent an outburst of disease. Quarantine,I believe, is a must for acclimatizing mostwildcaught cichlids. The “Corrienteras” got accli-matized very quickly and started to eat voraciouslyfrom the day of their arrival.

I keep my nebuliferum in a well circulated andoxygenated tank with a warm temperature of 28°’C.They seem to be doing just fine and show a healthyappetite. Of course, I pay much attention to water

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The location where P. nebuliferum was collected.

quality because, since I have observed the naturalconditions in which these fish live, you know theywould not settle for less.

I should remark the extraordinary personality ofthis fish, which does not get intimidated easily andhas a fantastic appetite. In this regard, I have notrouble feeding them. They take, willingly, frozenshrimp, fish meat, lettuce, spinach, brine shrimp aswell as live aquatic insects, which I collect near myhouse. I do, however, lean to a herbivorous diet. Itis hard to believe how much food this fish can take.

As in the wild, the Corrientera keeps in constantmovement in the aquarium as well. I provide themwith ceramic pots in the shape of caves that theyhappily use for cover and rest. Cichlid companywas, however, not easy to find because P. nebuli-ferum proved to be quite aggressive towards manyother cichlids that I tried to house them with. There-fore I decided to give them the whole 200 liters-tank to their own. I divided the tank, with plasticlight diffusers, into three compartments, one foreach individual.

Meanwhile the fry were eating like their adultcounterparts and growing steadily from one to aboutthree centimeters in two months. They were beingfed with live baby brine shrimp and softened spin-ach. Soon they started to show aggression amongthemselves, so I had to move them to larger quar-ters.

After a couple of months I noticed some kind ofattraction between two of the adult P. nebuliferum.I decided to remove the divider under close super-vision. Then the fish showed their magnificentbreeding dress which is hard to appreciate in theyellowish and dark water of their natural habitat.

The genital tubes were down the next morningand the pair spawned on the gravel. Nearly twohundred, small, yellowish, translucent eggs, aboutthree millimeters long, were deposited. During theentire length of the spawning, which took about twohours, the pair kept circling the spawning site. Af-ter a while, however, the eggs were eaten by thepair. I then had to put the divider in once again toprevent the pair blaming each other for their fail-ure, which is common with cichlasomines; the malewants to spawn again while the female holds nomore eggs.

Two weeks of heavy feeding passed before theywere ready to spawn again. I took the divider outand this time the pair chose the inside of the ce-ramic pot to spawn. This was more in agreement

with the natural situation where spawning takesplace in nests between the rocks in order to preventthe eggs from being washed away with the current.The second spawning, however, again did not suc-ceed, as the eggs were eaten the next morning. So Ihad to replace the divider once again.

Since then I have not placed the pair together,but I will surely do so again soon. This time theywill have to settle for breeding or I will have toremove the eggs from the parents to assure myselfof some fry.

New varieties of cichlids are being discoveredor re-discovered year after year. Central Americancichlids are gaining more popularity and many spe-cies, like P. nebuliferum, with an interesting col-oration and a great personality will indeed makepopular aquarium fish in the near future. Once theyare established in the hobby, the biology and habitsof the fish in the wild will begin to become impor-tant. With the growing interest, the availability ofnow rare cichlids will surely become a reality, as itsoon will be for P. nebuliferum.

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Introduction

In 1902 Vaillant and Pellegrin described for the firsttime the species “Cichlasoma” spinosissimum, albeitin the genus Heros. They described it from six speci-mens which were collected in the Rio Polochic inGuatemala. Two of the six specimens lacked the blackspots which were present in the other four. Pellegrin(1904) described these two as a variant of the nominalspecies and named it Cichlasoma spinosissimus var.immaculata.

In his famous revision of the genus Cichlasoma,Regan (1905) considered the difference mentionedbetween the two variants enough to regard “Cichla-soma” immaculatum as a valid species. After this lit-tle more was heard of this species.

In the beginning of the eighties a small cichlid ap-peared in the hobby which was initially identified asbeing “C.” spinosissimum. Both Staeck (1983) andMayland (1984) published in their books a photographof this cichlid which showed close affinities with “C.”spilurum. The authors Stawikowski and Werner (1985)duly note their doubts regarding the proper identifica-tion of this species and conclude that it is a geographi-cal variant of “C.” spilurum. The large distributionarea of this species indeed justifies the assumption that

A wildcaught “Cichlasoma” spinosissimum. Photo by Ton Woeltjes.

“Cichlasoma” spinosissimum (Vaillant & Pellegrin, 1902)Willem Heijns

several color variants exist.The question concerning the identity of “Cichla-

soma” spinosissimum was solved when, in 1989, somespecimens were collected in Guatemala and shippedto the Netherlands.

Taxonomy

Regan divided the genus Cichlasoma into severalgroups which he named sections. One of these sec-tions is Archocentrus, to which belong, besides “C.”spinosissimum and “C.” immaculatum, also cichlidslike “C.” nigrofasciatum, “C.” spilurum, “C.” sajicaand “C.” centrarchus. Within this section other group-ings can be made. The species “C.” spinosissimum,“C.” immaculatum and “C.” centrarchus consti-tute, in my opinion, a natural group which are moreclosely related to each other than to other members ofthe section Archocentrus.

A complication is created by the fact that “C.”centrarchus is regarded by Bussing (1976) as a sib-ling species of Herotilapia multispinosa. The latterspecies, however, does not belong to the genusCichlasoma let alone to the section Archocentrus. Theplacement of multispinosa in another genus is due tothe difference in the shape of its teeth compared with

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A pair of “Cichlasoma” centrarchus guarding their spawn. Photos by Willem Heijns.

Above: “Cichlasoma” spinosissimum, subadults.Below: Herotilapia multispinosa.

to those species currently in “Cichlasoma”. H.multispinosa has tricuspid teeth whereas all the spe-cies in “Cichlasoma” have uni- or bicuspid teeth.Nevertheless, I am of the opinion that these four spe-cies, “C.” spinosissimum, “C.” immaculatum, “C.”centrarchus and H. multispinosa, form a closely re-lated group where H. multispinosa might be a littlemore distant from the other three. Their close relation-ship is also expressed in their similar morphology andcolor patterns (see accompanying photographs).

If we consider the distribution patterns of the fourspecies we will immediately note that between the ar-eas of “C.” spinosissimum and “C.” immaculatum(Rio Polochic, Guatemala) and that of “C.” cen-trarchus and H. multispinosa (the great lakes of Nica-ragua) there is an enormous region where, up to now,no relatives of these species have been found. The factthat this area has not thoroughly been investigated,either by scientists or by aquarists might be a reasonfor this situation. I am sure that when this region isexplored in a more thorough fashion, new species andvariants are to be expected.

Ecology

“C.” spinosissimum inhabits calm waters with

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The distribution of “C.” spinosissimum (yellow), “C.” centrarchus (blue), andHerotilapia multispinosa (red).

plenty of aquatic weeds, which could be anticipatedregarding its close relationship with “C.” centrarchus.The specimens which were exported to the Nether-lands were collected in small tributaries of the Lagode Izabal, in the Rio Polochic drainage (the type lo-cality). They were found in almost stagnant water witha muddy bottom covered with a carpet of dead leaves.Their food consists mainly of insect-larvae which hidein the bottom and in between the rich growth of aquaticweeds. The pH of the water was around 6 and the hard-ness could not be measured. “C.” spinosissimum wasnot observed in the Lago de Izabal where the waterconditions noticeably differ from its tributaries. No re-cent collections have been made in the Rio Polochic,the most important water supply to the Lago de Izabal.“C.” immaculatum has not been found since its de-scription and it seems rather questionable whether twosuch closely related species could live sympatrically.Probably “C.” immaculatum turns out to be a geo-graphical variant after all.

“C.” spinosissimum seems to be a rather vulner-able cichlid. Several specimens died shortly after be-ing caught. Some others, which survived the long tripback home, acquired a certain type of fin rot to whichmost of them succumbed. Some individuals were curedafter a treatment with tetracycline.

In the aquarium

At this moment experience with “C.” spinosissimumin captivity is rather scarce. The few specimens thatmade it to the Netherlands were bred but they did not

survive after all. The observa-tion of one breeder was that“C.” spinosissimum is a veryshy aquarium fish, whichhides among the rockwork ofthe tank. His pair spawned in-side a cave which might leadto the conclusion that it is acave-brooder. This, however,is unlikely as its close relativesspawn on open substrates.Like “C.” centrarchus and H.multispinosa it “hangs” thelarvae on the roots of plants oragainst the side of the aqua-rium.

My first experience withthis cichlid started with 30 ju-veniles, barely in the free-swimming state. I fed them

large amounts of Artemia and encountered the firstproblems when I changed to another type of food inorder to cope with their needs. Although the food thatI offered, such as Cyclops and mosquito larvae, wascertainly small enough to be eaten, they only reluc-tantly considered it as food. Their growth stunted andsome died. At one time too many dead bodies mademe think a disease had played havoc with the fish.Only three fish were left over when a normal situationreturned. At this moment they are about 8 cm in sizeand I think I have two males and one female. One ofthe males is clearly the boss and chases the others fromits territory. These two, however, show now and againsome courting behavior. I have patience.....

References

BUSSING, W. A. (1976) Geographic distribution of theSan Juan ichthyofauna of Central America with re-marks on its origin and ecology. In: Investigationsof the ichthyofauna of Nicaraguan lakes. T. B.Thorson (Ed.), Lincoln, Nebraska.

MAYLAND, H. J. (1984) Mittelamerika. Cichliden undLebendgebärende. Landbuch Verlag, Hannover.

STAECK, W. (1983) Cichliden — Entdeckungen undNeuimporte. Band III. Engelbert Pfriem Verlag,Wuppertal.

STAWIKOWSKI, R. & U. WERNER (1985) Die Buntbarscheder neuen Welt. Mittelamerika. Kernen Verlag, Es-sen.

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When Carl Hubbs described “Cichlasoma”’ synspilum,in 1935, he pointed to a close relationship of this spe-cies with Heros melanurus, which had been describedby Albert Günther in 1862. The status of “C.”synspilum had been in question for a long time andmany authors believed that it should be regarded as asynonym to “C.” melanurum. “C.” synspilum wasintroduced into the hobby long ago but the locality ofthe origin of the red variety, which has established it-self firmly among hobbyists, is still unknown. Theholotype of this species, however, was collected in theRio San Pedro de Martir, a tributary of the RioUsumacinta. This location is just 100 km away fromthe type locality of “C.” melanurum, which had beendescribed from specimens collected in Lago Petén. Itcould be the proximity of both type localities that mayhave misled some authors who regard “C.” synspilumas a synonym. Although the distance between the twotype localities is small, these two waters belong to twodifferent drainages. This may be the main argument forthose authors who regard both cichlids as valid species.

Hubbs found that the black, horizontal band, whichruns on the caudal peduncle towards the center of thebody, marks the difference between the two species.In “C.” melanurum this band runs almost mid-lateral

A female “Cichlasoma” melanurum in breeding coloration. Photo by Willem Heijns.

Distribution of “C.” melanurum (red) and “C.” synspilum (blue).

“Cichlasoma” melanurum (Günther, 1862)

on the peduncle whereas it “dips” down and runs closerto the anal fin in “C.” synspilum.

Another usable characteristic of “C.” melanurumis the golden-yellow coloration in adult specimens. Thespecimen in the photograph above was caught in LagoPetén, in 1989, and represents the first “C.” melanurumthat had been exported to the Netherlands. It is notcertain if different color variants exist of this specieswhich are known for “C.” synspilum.

“C.” melanurum is, like its close relative, a veg-etarian but likes any type of aquarium fare given.

Willem Heijns

72

The species of the section Herichthys in the genus“Cichlasoma” belong to the group of most north-erly distributed cichlids in America. One of them isthe Texas cichlid. The Herichthys cichlids aremainly found in the Atlantic drainage of México.In the Pacific drainage “C.” beani, a member ofthe Parapetenia section, inhabits an even morenortherly region but this still lies in México.

The most important group in Herichthys is, with-out doubt, the species closely related to the Texascichlid, “C.” cyanoguttatum, to which also “C.”carpinte belongs. The distribution of this groupranges from the Rio Grande and its tributaries inTexas to the drainage of the Rio Pánuco in the Mexi-can state San Luis Potosí. The entire distributionlies on the Atlantic drainage of Central America.The description of a Herichthys found in the Pa-cific drainage (Hernandez Rolon, Revue fr. Cichlid.,Sept. 1990) must be an error which may have beencaused by an introduction of an Herichthys by menin this area.

During a recent trip through the drainage area ofthe Rio Pánuco we decided to travel towards thesouth in order to find some river systems which areindependent from the Rio Pánuco and which mightbe inhabited by Herichthys as well, preferably dif-

A wildcaught male “Cichlasoma” sp. “Pantepec”. Photos by Willem Heijns

The Rio Pantepec where the new Herichthys was found.

“Cichlasoma” sp. “Pantepec”

ferent ones to those found in the Pánuco system.We traveled from Tampico towards Vera Cruz andat about 150 km south of Tampico we came to thesmall town of Alamo where the road crossed, via avery long bridge, the Rio Pantepec. Assisted by afew local boys we collected a new Herichthys,which we immediately named Turquoise Herichthysbecause of its coloration (see photograph). In col-oration it seems to be intermediate between theTamasopo Herichthys (Rio Pánuco) and the vari-ant/species which had been collected some yearsago near Nautla (Vera Cruz).

Willem Heijns

73

About ten miles south of the crossing of the RioPantepec, where we had collected the TurquoiseHerichthys, we stumbled on the town of Poza Rica.In the center of the town a bridge crosses the RioCazones which invited us to throw our nets in searchof more cichlids. Probably due to the rather fastflowing current we could not catch any fish and wedecided to look for a calmer side-stream to try ourluck again. Searching for such a stream we arrivedon the outskirts of the town where we found a smellypool which was probably used as a sewer by thesurrounding houses. While one of the housewiveskept an eye on our car and valuables, we wet ourfeet in the far from inviting pool. Contrary to ourexpectations we collected some beautiful cichlids.The pool was part of a stream that looked cleanerdownstream; so we decided to continue catching alittle further downstream. It proved quite difficultto collect enough specimens as the populationseemed rather thinly spread.

“C.” sp. “Poza Rica” resembles in many respectsthe Turquoise Herichthys from the Rio Pantepec butcan be distinguished by the red coloration, espe-cially on the cheek and gillcover.

(At present it is difficult to assess the status ofall the Herichthys found in the different streams of

A wildcaught male “Cichlasoma” sp. “Poza Rica”. Photos by Willem Heijns

“Cichlasoma” sp. “Poza Rica”, a female.

“Cichlasoma” sp. “Poza Rica”

México. It seems, though, that a good part of themcan be classified as geographical variants of “C.”carpinte. The more populations that are investi-gated, the better we will understand the variationamong the species of the Herichthys section of thegenus “Cichlasoma” . Geographically-relatedchanges in coloration are a common feature incichlids and should not lead to a recognition of asmany Herichthys species as there are streams inMéxico. Ed.)

Willem Heijns

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The distribution of “Cichlasoma” bartoni is restrictedto the upper Rio Verde. The largest population is foundin the Media Luna, a spring in the province of SanLuis Potosí, México. There are some smallerpopulations north of the Media Luna. They are mostlyfound in spring-fed streams and rivers. I have observedonly the population in the Media Luna. There, “C.”bartoni is probably the most abundant cichlid. Unfor-tunately its position is slowly taken over by “C.”carpinte which was introduced from the Rio Verde. In1980 a canal from the spring – built to irrigate the land– was completed. This canal, however, lies below thefloodplain of the Rio Verde. In the rainy season theriver occasionally spills into the canal and thereby in-troduces its fish-fauna into the Media Luna system.

“C.” bartoni is a rather small member of theParapetenia section in the genus “Cichlasoma”. Themaximum recorded size is 20 cm but the majority ofthe adult, breeding individuals measure between 9 and14 cm. Almost any individual over 10 cm in size showsits breeding dress. “C.” bartoni is probably one of thefew cichlids in the northern half of México whichbreeds throughout the year. The temperature of thewater in the Media Luna is rather constant and readsabout 25° C throughout the year. The water tempera-ture further away from the feedingspring fluctuates

A pair “Cichlasoma” bartoni with fry, photographed in the Media Luna.

“Cichlasoma” bartoni (Bean, 1892)

much more with the seasons. In late summer, the warmspring water – with a hardness of more than 50 de-grees – is diluted with the colder rain water. Most cich-lids in the Rio Pánuco drainage breed at the beginningof the year, in the dry season, because the water has itshighest temperature. “C.” bartoni is normally foundclose to the origin of the spring. In Media Luna, thespring produces a substantial flow of water in which itis difficult to move about. Nevertheless, many breed-ing pairs are found in the strong current. To preventtheir broods from drifting away they dig pits in thesand, preferably behind a small rock. The turbulence,caused by the rock, brings drifting food particles rightinto the nest. Spawns may number more then 200 fry.Breeding pairs are found one to two meters apart.

“C.” bartoni is a predator. It searches the bottom ofthe stream for something edible, usually invertebrates.Sometimes juvenile’”C.” bartoni’ profit from the for-aging method of “C.” labridens with which it livessympatrically (see photo next page). Despite the mul-titude of bartoni fry in the spring only a few semi-adult individuals, which are solitary, are observed.

Ad Konings

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In captivity “Cichlasoma” labridens attains a maxi-mum total length of approximately 25 cm but mostadult individuals seen in their natural environment havea length between 12 and 16 cm. “C.” labridens has arather wide distribution and is therefore found in dif-ferently colored geographical races. It is found insprings, streams and rivers in the catchment area ofthe Rio Pánuco. The yellow race – of which a pair isshown in the picture – is known only from Media Lunaand the upper Rio Verde. Most other populations havea white breeding coloration. The neutral color-pattern,however, differs marginally among the differentpopulations.

In the Media Luna, “C.” labridens was originallyfound sympatrically only with “C.” bartoni, but theirhabitat is now threatened by introduced tilapiines and,especially for “C.” labridens, by “C.” carpinte. Likethe latter “C.” labridens’ searches for food on the bot-tom of the stream and both have the same, peculiarway of doing so. By wagging the entire body the sandis cleared from the debris. The uncovered invertebrates,usually small snails, are then collected by the fish. Both“C.” labridens and “C.” carpinte feed on snails sincethe tilapiines eat most of the edible plants. “C.”carpinte is known to eat plants as well but thanks to

A pair “Cichlasoma” labridens leading their fry through the Media Luna.

A “C.” labridens wags the sand clear of debris while juvenile“C.” bartoni pick their choice from the shower of mulch.

“Cichlasoma” labridens (Pellegrin, 1903)

the introduced tilapiines it competes effectively with“C.” labridens for the hidden invertebrates.

“C.” labridens usually spawns in spring. The eggsare frequently deposited on the stem or exposed rootsof waterplants – there are many Nymphea in theMéxican waters. When the fry have reached the free-swiming stage they are led by the parents through thehabitat. From time to time one of the parents clearsthe substrate by wagging and all the youngsters willdive in the clouds of debris to look for something ed-ible.

Ad Konings

76

It is 6.30 am. The weak outboard motor slices, withnoticeable effort, the large canoe through the seem-ingly ropy water of the Yarina Cocha. The water iscompletely black and the thin shreds of mist that waftover the surface give the scene something mystical.Frequently, the black-mirroring surface is broken by agroup of breathtaking dolphins. The rising sun lets usfeel what incredible high temperatures we will haveto endure again this day while parrots flying over our

A wildcaught male Tahuantinsuyoa macantzatza from the Rio Huacamayo. Photos by Ron Bernhard.

SOUTH AMERICANCICHLIDS

Tahuantinsuyoa macantzatza Kullander, 1986

Ron Bernhard

heads scream their loud protest. Some moments laterwe anchor at the fishermen’s village Yarina from wherewe will continue our trip by car.

This was the start of a collecting trip to the interiorof Peru. After a rather troublesome flight from Limato Pucallpa we (myself and eleven other aquarists) wereawaited by the staff of La Cabana, the lodge which wewould use as a basis for our trips. A little later we wereon our way on our first, nightly trip on the warm, pro-

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Tahuantinsuyoa macantzatza with jaws extended.

lific waters of Yarina Cocha.In the coming weeks we would make several expe-

ditions in this area which abounds with rivers andstreams. One of these trips supplied the introductionto this article. We were the four of us and had decidedto drive towards the Andean mountains, to the typelocality of Aequidens patricki. It proved rather diffi-cult to rent a taxi in Yarina to drive us to Aquaytia. Notthat the distance was a problem but the fact that thetype locality of this pretty cichlid, Aquaytia, lies in themiddle of Peru’s largest “coke” plantatations. Finallywe were on our way, much later than planned.

After we had located and collected A. patricki we re-turned to the lodge. Notwithstanding the fact that ourexpedition resulted in 22 specimens, Ulrich Minde wasdetermined to repeat the procedure the following day.

When we drove to Aquaytia our road had crossed asmall river, the Rio Huacamayo. Due to our late de-parture we did not have time to collect in this river butUlrich, who went on his own, had. The water in theriver was clear and there was a weak current. In hisnet were caught some cichlids with a strong resem-blance to species of Bujurquina (the larvophilousmouthbrooders which previously belonged to Aequi-dens). He brought some specimens to the lodge butmost of them proved to be sensitive and were subse-quently lost during transport. The ones that had sur-vived the trip died during the following days. Fortu-nately there was one female with almost matured lar-vae. These were treated with great care and survivedthe trip back home.

After Ulrich had raised the fry to a reasonable sizehe gave me seven. Meanwhile we had figured out theidentity of this little cichlid. Its resemblance toBujurquina species was striking but it had one featurewhich was not shared with these cichlids. All speciesin Bujurquina have a band between the eyes whichlies forward on the head. In our species the band startedat the eye but ran backwards, towards the dorsal fin.This seemingly simple distinction was enough to iden-tify our cichlid, using Kullander’s book “The cichlidfishes of the Amazon river drainage of Peru”, as theInca Stonefish or Tahuantinsuyoa macantzatza.

The common name is derived from the fact that thebottom of all rivers inhabited by T. macantzatza con-sists of pebbles and rocks. T. macantzatza is a slenderlybuilt cichlid with a rather heavy head with thick lips,and is laterally somewhat compressed. The unpairedfins are, in both sexes, rounded. There are six verticalbars on the body which become distinct when the fish isexcited. The maximum size of T. macantzatza is about

13’cm for the male; the female remains a little smaller.The difference between the sexes is rather small.

The coloration of the male is a bit more intensive andhis proportions are a little more robust than those ofthe female. During the breeding period the female hasa vague dark patch in the dorsal fin.

The behavior of the Inca Stonefish preceding thespawning, was rather strange. At one moment both fishwere close together supposedly preparing for spawn-ing and some hours later the female was chased to everycorner of the tank. This kind of ritual lasted for abouttwo weeks after which the pair-formation was a fact.Both male and female were frequently found side byside with head down and all fins erect while the entirebody quivered.

The eggs were deposited on a small chip of wood.There was barely enough room for them. I recalled thisbehavior from Bujurquina vittata, who, when I was pho-tographing them, took the chip of wood with the eggsand carried it out of reach of my camera.

Unfortunately, I have never succeeded in successfullybreeding T. macantzatza. Every time the female swal-lowed the eggs two or three days after she had takenthem into her mouth. Uwe Werner, who has spawnedthis species, found that the female took the larvae in hermouth after 52 hours (pH of 7.5 and 26°’C). Also themale participated in the care of the larvae. He sometimestook the larvae from the female while she fed.

The spawns are rather small; about 40 to 50 eggswere deposited each time. As soon as the fry are bigenough to forage on their own they are released by thefemale. The first few days after the release she occa-sionally takes them back into her mouth but soon thefry go their own way.

T. macantzatza is a very interesting cichlid whichneeds a medium size aquarium. Regrettably, a widedistribution among hobbyists cannot be expected soonas it has only sporadically been bred in captivity.

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For many years Biotodoma cupido seems to have been afrequently kept cichlid. It is mentioned in most aquariumliterature but in reality it is difficult to obtain just a fewspecimens. The best chance one has is to looka for “con-taminants” in shipments of Geophagus surinamensis orSatanoperca leucosticta. Another species of the genusBiotodoma, B. wavrini, is regularly exported but onlyduring the last few years.

Biotodoma sp. “Santarém” was exported for the firsttime in 1988. A German importer in Munich offered thisspecies as “the most beautiful Biotodoma”. The accom-panying photograph is a testimony to this title.

Biotodoma sp. “Santarém” is found near the town withthe same name (in Brazil) in an enormous swamp wherethe Rio Tapajós flows into the Amazon. The water levelin this swampy delta varies dramatically with the sea-sons. It can be 15 meters higher at the end of the rainyseason than before the rains started. An unknown lake inthis vast area is used as the collection site by the ex-porter. The collection of aquarium fishes in this area ispossible only during the dry season, from August to De-cember. If something pops up it can happen that this spe-cies is not available for more than a year.

Like the other species in the genus, Biotodoma sp.“Santarém” grows to a maximum size of approximately

A fully mature male Biotodoma sp. “Santarém”. Photos by Ron Bernhard.

Biotodoma sp. “Santarem”

Ron Bernhard

12 cm. It is a very placid cichlid, which should be kept inan aquarium with ample hiding places. In particular drift-wood and some larger aquatic plants are essential for asuccessful spawning.

The male is recognized by the filamentous extensionsof the unpaired fins and its splendid coloration. From asize of about 6 cm individuals can also be sexed by ex-amining the vents. The variation in the coloration of themales is remarkable. In a single shipment of wildcaughtspecimens, some males have a completely colored tailfin whereas others only have a white upper and loweredge. Adult males acquire a fantastic finnage; dependingon the light the dorsal fin can be orange and green or redwith blue. The exceptional finnage, as the male in thephotograph shows, can be attained only in a tank with-out any dominant or aggressive inhabitants. The best wayto keep this species is pairwise, possibly accompaniedby small tetras or catfish.

Breeding B. sp. “Santarém” is, in comparison to otherspecies of the genus, relative simple. Although I havekept, on several occasions, B. wavrini and B. cupido (fromnortherly regions of South America), it was now for thefirst time that a species of Biotodoma spawned in myaquarium.

The first specimens I could obtain were placed in a

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A female Biotodoma sp. “Santarém” guarding the eggs.

130 cm-long aquarium. Already after one day a pair hadformed and the male had dug a pit. The pair harmonizedwell and I expected them to spawn soon. But two weekslater I found the female between the plants and she seemedto have lost her colors. The new color pattern consistedof a white dorsal fin, the body and head rather dark ex-cept for the ventral region which was white again. Theblue-fluorescing streaks, which were usually seen on thehead, were diminished to vague blue spots. The initialthought that the pair had quarrelled soon vanished whenI noticed, below the female, a few hundred eggs sway-ing in the stream caused by the fanning of her pectoralfins. The eggs were attached to the substrate by just athin thread. After two days my joy disappeared becausethe female had eaten the eggs. Later spawnings failed aswell; this was probably due to the high pH and conduc-tivity of my tap water. Up to now Biotodoma had be-haved as a purely monogamous cichlid. A new breedingsetup cast some doubt on this assumption.

A new group, consisting of one male and three fe-males, was introduced into a 90 cm-long breeding tank.This time I had adjusted the water to a pH of 4.5 and aconductivity of 80 microSiemens (at 25° C). As soon asthe fish were released the females chose each their ownlittle territory. The male regarded the entire tank as its

property. After one week he started courting the females,all three of them. The complete change in coloration in afemale indicated that there were eggs. The next day, how-ever, I noticed all three females with a breeding colora-tion! All three females had spawned with the same malewithin 48 hours. The male moved from one female to theother and seemed to be at his wit’s end regarding thisunusual situation. One day later the male had made itsdecision and stayed with one female. Together theychased the other females away from their spawn. In theend only the eggs of the pair hatched and the remainingtwo females had to accomodate themselves in a cornerof the aquarium. Just five days after hatching the femaleled the fry through the tank.

The water quality seemed to be of crucial importancein breeding this Biotodoma. After its polygamous ex-cesses, the male restricted itself to one female only. Thepolygamous behavior observed might thus have beencaused by a coincidental presence of three ripe females.

The fry were easily raised. They grew fast and weresoon spread among other hobbyists. I hope they have asmuch pleasure and luck with this cichlid as I had, be-cause Biotodma sp. “Santarém” is a jewel among thecichlids, in coloration as well as in behavior.

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All representatives of the genus Retroculus were,until recently, catagorized as cichlids whichaquarists had never seen alive. The only illustra-tion I had ever seen was a colored drawing of R.lapidifer. The drawing accompanied a scientificdescription of Chromys lapidifera which DeCastelnau published in 1855. The illustration im-pressed me very much as I could hardly imagine aSouth American cichlid which would look like thisspecies.

For years I wondered how this Retroculus wouldlook like alive and it came as a surprise when I reada small article in DATZ (Vol. 40; Nr. 12) about thefirst import of this cichlid. Shortly afterwards it wasseen in the trade.

When De Castelnau described Chromys lapidi-fera he could barely guess that, before the end ofthe century, it would change its generic name threetimes. Finally, Regan (1906) placed it in the stillaccepted genus Retroculus. Besides R. lapidifer twoother species have also been described in this ge-nus, namely R. xinguensis and R. septentrionalis.

R. lapidifer is distributed in the drainages of theRio Tocantíns, Rio Araquaia and Rio Guamá, in-cluding their tributaries. It is known that severalgeographical variants exist. The specimens shownin the photographs were collected in the Rio Guamá.They lack the vertical striping in the tail fin and thedorsal fin is almost without markings. The throatand upper lip are yellow and there are some red spotson the ventral side.

The variety from the Rio Tocantíns, which wasexported to Europe recently, has a lot more colors.The dorsal fin shows more red and white, while theupper lip is blue. There are many red spots on thebelly.

In the aquarium R. lapidifer behaves as an idealaquarium fish. It eats any type of aquarium fare apartfrom vegetable matter. Even small tetras, which areleft unharmed by species of e.g Satanoperca, areconsidered as food. Its favorite occupation, in whichit reveals the close relationship with Geophagus, isto poke its snout up to the eyes in the sand, longafter it has been fed. The way it dives into thesubstrate is rather unique. Before it “strikes”, it putsitself in an almost vertical position. A quick dartforwards plunges the head in the sand. The eyes areplaced far back, away from the mouth, a feature thatgave these species the generic name Retroculus.

The sexes are difficult to identify. It seems thatmales have thicker lips and a more intensive col-

Retroculus lapidifer (De Castelnau, 1855)

Ron Bernhard

oration than females. I also noticed that the shapeof the female’s head is less robust, as in many spe-cies of Satanoperca. The female’s snout seems tobe shorter and her head, as a whole, looks more“friendly”.

Adult R. lapidifer reach a size over 20 cm; this,together with its sometimes temperamental behaviorindicates that it needs a large aquarium.

Already in the description of De Castelnau, in1855, the breeding behavior of Chromys lapidiferawas discussed. He wrote that it makes a nest of smallpebbles, which are arranged in a round heap. Thisis confirmed by recent observations in the wild.Moreover some successful spawnings have beenrecorded in captivity. Hartwin Kiefel reported hisobservations on the breeding pair in his aquarium.The eggs were deposited on a flat stone and wereimmediately covered by small pebbles and stones.These were collected from every corner in the tankand thrown on the spawn! This behavior gave thecichlid its scientific name; lapidifer means “stone-carrier”.

Unfortunately the eggs never hatched, althoughthe pair spawned several times. This is probably dueto the differences in the water chemistry and maybethe oxygen content of the water. The pH of the wa-ter in its natural habitat measured barely above 7while the hardness was not measurable.

Retroculus species prefer fast flowing waters andbelong to the rheophilic cichlids. Like Steatocranusand Teleogramma, they have lost the ability to hovermotionlessly in the water. Their buoyancy compen-sating organ, the swimbladder, is only rudimentarilypresent. When R. lapidifer swims, it gives the im-pression as if something is pulling it down. We mustnot forget that this cichlid lives in waters that flowtoo fast for us to stand upright. The decoration andfiltration of the aquarium require some extra atten-tion. The bottom should be covered with coarsegravel or medium sized pebbles. The water must bein motion al the time; a powerful power filter shouldcreate a visible current in the tank. R. lapidifer likesto watch its environment from an elevated position.Some flat stones, slabs or flagstone, will make iteasy for them. They also use these as spawning-sites.

If the tank has been prepared in the way men-tioned, one will enjoy the presence of a couple ofR. lapidifer even if a successful spawning is not tobe expected for some time.

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Retroculus lapidifer from the Rio Guamá.

Retroculus lapidifer is a very attractive cichlid. Photos by Ron Bernhard.

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Most Crenicichla rarely exceed a size of more than25 cm in their natural habitat, but there is a groupamong the pike cichlids which normally grow over30 cm in length. Many common features like mor-phology, meristics of fins and scales, and similari-ties in the development of fry to adult fish, indicatethat this group of large pike cichlids may form adistinct section within the genus Crenicichla. Al-though imports of such large Crenicichla have beenrare, at least one species of this group was infre-quently shipped to Europe in the last ten years. Thisspecies, which was collected in Venuzuela, becameknown under the name C. strigata. The namestrigata is derived from the fact that juveniles havea pattern consisting of horizontal lines on the body.Besides these lines there are also black spots on thehead, arranged in an attractive pattern. A drawingof a similarly patterned cichlid in Regan’s Creni-cichla revision (1905) strongly suggested that theVenezuelan pike cichlid was conspecific with C.strigata Günther, 1862. The fact that there are atleast three valid species with this type of juvenilecolor pattern was not known until recently.

The first doubts were cast on the identity of theVenezuelan species when, about two years ago, a“new” Crenicichla was found in the Rio Capim

An adult Crenicichla strigata from the Rio Capim. Photos by Frank Warzel.

Some large Crenicichla

Frank Warzel

drainage. The Rio Capim was mentioned, in 1862,as one of the two original localities of C. strigata.The “new” pike cichlid closely resembled the Ven-ezuelan species, but had a greenish coloration anda conspicuous white-edged blotch behind thegillcover. In the same shipment of these greenCrenicichla there were several, more than 20 cm-long, pikes which showed an attractive pattern ofspots and stripes. The pattern was identical with thatin the drawing of C. strigata in Regan’s publica-tion of 1905. Later, when the green Crenicichlagained a similar size to that of the striped individu-als, it became clear that both belonged to the samespecies. The obvious conclusion that this species isthe true C. strigata is not basically wrong but stillneeds some scientific verification. For certain, thespecies from the Capim drainage (and also a greenCrenicichla from the Rio Tocantins) is notconspecific with the aquaristically known “Strigata”from Venezuela.

Another species, with Manaus as its alleged ori-gin, likewise shares similarities with the former“Strigata” from Venezuela. It has, however, a moreelongated body shape and caudal peduncle andshows a narrow black and white band in the dorsalfin. Moreover, the blotch on the caudal peduncle

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A female Crenicichla lugubris allegedly from Manaus.

has an irregular shape without a light border, whichis seen in similar-sized Venezuelan pike cichlids.

A cichlid with a great resemblance to this spe-cies, Crenicichla lugubris, was described by Heckelin 1840. In Heckel’s description there is no men-tion of a light band in the dorsal, but this is a typi-cal characteristic of females which is regularly ob-served in the other species of this group. Accordingto recent opinion (Kullander & Nijssen, 1989) C.lugubris is found only in the Corantijn system, atthe border between Surinam and British Guyana,in the Essequibo river, in the Rio Branco and in thelower Rio Negro. Besides morphological conformi-ties, the alleged distribution of the species (Manaus)further suggests that we are dealing here with thetrue C. lugubris.

The coloration pattern, as far as one can regard itas such, consists of a plain, light gray on the bodyand some red pigment in the ventral fins. Duringthe breeding period, which usually takes place inwinter, the fish changes its garment completely andone would think it an entirely different cichlid. Thelower part of the head becomes yellow and orange,the ventral fins become bright red, while dark, vio-let zones are visible on the flank. As in mostCrenicichla, the ventral region of the female be-

comes lighter with some reddish hues. Unfortu-nately nothing is known, aquaristically speaking,about the male coloration or about that of juvenilessince only a single female has been exported.

Something more is known about Crenicichlamarmorata Pellegrin, 1904, which was importedfrom the lower Rio Tapajós. This species, which isreadily identified when adult, also shows a juve-nile pattern of stripes and spots and is in this phasehardly distinguishable from the other species. Notuntil the moment of the color-change was the au-thor convinced he had kept semi-adult C. strigata.In retrospect, the horizontal stripe of juvenile C.marmorata did not seem to be so sharply borderedas it is in C. strigata.

Although juveniles all look alike, adult C.marmorata show an individual color pattern. Evenfish from the same spawn display a variable pat-tern. While some individuals have black blotchesalternating with colored ones on the dorsal part ofthe body, others have a horizontal row of coloredspots. The most common pattern is a combinationof these two basic patterns.

Besides that from Santarém on the lower RioTapajós more populations of C. marmorata areknown. The Dutch ichthyologist Alex Ploeg (1987)

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A juvenile Crenicichla marmorata. Photo by F. Warzel.

A juvenile C. sp. “Strigata Venezuela”. Photo by F. Warzel.

An adult male Crenicichla marmorata from Santarém. Photo by Frank Warzel.

A juvenile C. strigata. Photo by Rainer Harnoß.

An adult C. sp. “Strigata Venezuela”. Photo by F. Warzel.

85

describes this species from the lower Rio Trombetas,where the population has nine irregular bars thatreach onto the ventral region. Three specimens fromthe Rio Madeira system show, in contrast to the pre-vious population, a more dorsally oriented and par-tially connected blotch pattern. The head patternsare different, too. While the Trombetas marmoratahas a completely black head, except for a small zonebehind the eye, Ploeg found the head of the Ma-deira marmorata dotted with small, black spots.

It will probably take years before all systematicquestions regarding Crenicichla are solved.

Less problematic is the maintenance of thesecichlids, although some experience with large fishis required (Stawikowski & Werner, 1988).

One of the peculiarities of these large species isthe social behavior of the juveniles, who, when theystill have the characteristic spots-and-stripes pat-tern, are very peaceful and seek contact with eachother. The cohesion of a group can be so strong thatthey jointly chase intruders from their territory. Inthe natural habitat this may be the duty of the par-ents, who guard their offspring for a period of sixmonths or longer. After about one year – the periodmay vary according to species and conditions – thechange in color takes place. A concomitant changein their behavior is noticeable, and skirmishesamong the members of the group are frequentlyseen. In tanks larger than 150 cm these rebelliouspikes can still be kept together, provided that theyequal each other in size.

Like most large cichlids these pike cichlids aremonogamous, i.e. they breed in pairs. In the re-stricted quarters of an aquarium this might some-times lead to quarrels between the pair. The dis-tinction between the sexes is rather difficult as theregularly observed extensions of the unpaired finsin males of other species are not seen in Crenicichla.In general, the females have, however, a light bandor zone in the upper part of the dorsal fin.

Regrettably, a successful spawning, in captivity,of one of these large Crenicichla has not yet takenplace, although courting behavior and females withripe eggs have been observed. Probably certain en-vironmental factors, which can only partially becreated in an aquarium, play a crucial role here. Itwould be a shame if, in future, we are not able tobreed these cichlids, because then we must continueto depend entirely on imports of these magnificentfish. Many of them will then probably “ooze away”in one tank or another.

References

GÜNTHER, A. (1862) Catalogue of the fishes in the Brit-ish Museum. Vol. IV. London.

HECKEL, J. (1840) Johann Natterer’s neue Flussfischebrasiliens nach den Beobachtungen und Mit-teilungen des Entdeckers beschrieben. Ann. wien.Mus. Nat. Vol. 2; pp 327-470.

KULLANDER, S. O. & NIJSSEN, H. (1989) The cichlids ofSurinam. Leiden.

PELLEGRIN, J. (1904) Contribution à l’étude anatomique,biologique et taxonomique des poissons de la familledes Cichlides. Mém. Soc. zool. France, 16; pp 41-399.

PLOEG, A. (1987) Crenicichla marmorata Pellegrin,1904 du bassin du Rio Trombetas, Brésil, nouvelledescription illustrée. Rev. fr. Aquariol., 14 (3); pp85-88.

REGAN, C. T. (1905) A revision of the fishes of theSouth American cichlid genera Crenacara, Batra-chops and Crenicichla. Proc. zool. Soc. London. pp152-168.

STAWIKOWSKI, R. & U. WERNER (1988) Die Buntbarscheder neuen Welt—Südamerika. Essen.

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Normally one would not expect to find a small andattractively colored cichlid among the species of agenus which has the reputation of consisting oflarge, unattractive predators. However, the genusCrenicichla includes a group of small (in compari-son to the other species) dwarf-sized cichlids whichattain, under natural circumstances, a maximumlength of less than 10 cm. Not only the size but alsothe elegant and graceful appearance and the inter-esting courting behavior of these dwarf Crenicichlasmake them very rewarding aquarium residents.

Until recently only two species of this group wereinfrequently imported. In 1990 a third species,Crenicichla compressiceps, was imported for thefirst time. The “new” dwarf Crenicichla shows sur-prisingly little similarity to the other species of thegroup. This is mainly due to the difference inbiotopes which these species inhabit. Whereas C.regani and C. notophthalmus are found mainly inthe weedy, shallow regions near riverbanks, C.compressiceps is observed only among the rubbleand rocks of the lower Tocantíns River system.When I observed C. compressiceps in their habitat,I noted that they were not at all shy and they rarely

A wildcaught male Crenicichla compressiceps. Photo by Frank Warzel.

Crenicichla compressiceps Ploeg, 1986

hid under rocks. Large caves and gaps between therocks were principally avoided. This remarkablebehavior might be explained by the fact that twomuch larger Crenicichla species inhabit the samebiotope and they commonly use these caves as shel-ter. The agile behavior of C. compressiceps mightalso be related to the situation in their home wa-ters. This dwarf Crenicichla has a lot of tempera-ment and males require rather large territories fromwhich they energetically chase rival males.

A suitable aquarium will be larger than 80 cmand have ample rockwork. Hiding places should becreated in the upper part of smaller tanks. Victim-ised fish will thus find an easy refuge. In return forits aggressive inclination we are rewarded with anattractive coloration.

According to the experiences gathered from keep-ing this species in captivity we should not expectmany difficulties. C. compressiceps relishes anytype of frozen and live food. We must be carefulnot to overfeed them because their enthusiasm forfood may get the better of them.

Frank Warzel

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In 1986 the Dutch ichthyologist Alex Ploeg describedsome new Crenicichla species, which were collectedin the beginning of the eighties during several scien-tific expeditions. These expeditions were undertakento survey the local fish fauna, as a construction of anenormous dam would soon for ever destroy the rapidshabitat in the lower Tocantíns.

Our collection trip in late summer 1990 brought usto Marabá, a small town upstream of the now over170 km (!) long reservoir. One of the rheophilic spe-cies known from this region is Crenicichla cyclostoma.The name means “round mouth” and is derived fromthe fact that, when viewed from above, the lower lip(which is larger than the upper) is almost perfectlyround. As in the sympatric C. compressiceps, the headand body of C. cyclostoma are laterally compressed,which is a remarkable feature considering the wide-mouthed appearance of most other species of the so-called Batrachops group. The maximum size of C.cyclostoma is about 13’cm, which is again in contrastto that of its closest relatives. The large teeth on theouter jaws (sometimes also seen when mouth is closed)are another characteristic of this peculiar species.

C. cyclostoma, which lives a hidden life under therocks of its biotope, adapted gradually to its new en-

The head of Crenicichla cyclostoma; note the large teeth. Photos by Frank Warzel.

Crenicichla cyclostoma.

Crenicichla cyclostoma Ploeg, 1986

vironment and greedily accepted frozen foods.In the wild the diet probably consists of insect lar-

vae and small crustaceans. For maintenance the wateris of minor importance but enough shelter, in form ofrocks, should be created.

C. cyclostoma has the ability to change its colorpattern rapidly, especially upon confrontation withconspecifics. The dominant individual acquires a lightbody coloration with vertical bands whereas theweaker one becomes dark gray with a light gray, hori-zontal line. During the breeding period females havea bright red band in the trailing part of the dorsal fin.

Frank Warzel

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Symphysodon aequifasciatus Pellegrin, 1904

A very decorative pair of a strain of Symphysodon aequifasciatus.

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Symphysodon aequifasciatus Pellegrin, 1904

A beautiful female of a red-turquoise strain with a regular pattern

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It is advisable to offer most cichlid species ample hidingplaces in the aquarium for the available space is usuallynot sufficient to let all inhabitants coexist harmlessly to-gether. Shelter in tanks is normally created by placingrocks and other heavy artifacts in such a position that theinhabitant can find refuge among them. A second reasonto place heavy rocks in your tank is to imitate the naturalbiotope of the cichlids and to increase the surface areaonto which algae, the natural food of many rock-dwell-ing cichlids, can settle. From an aesthetic point of viewlarge rocks are recommended as they will give the im-pression of a larger tank. Because of the fact that mostaquarium bottoms are made of glass, large rocks are haz-ardous. To abridge the wish of an aesthetically approvedreef and that of an intact glass bottom, the followingmethod describes how one can make featherweight rocks.

What we need is a rock, which serves as an example,gypsum, clay and a wooden box, which we custom makefor the size of the rock.

Casting the mold

We construct a box from wood, which fits the rock byat least 3 cm on all sides. On the bottom we place a ringof clay, at least 3 cm high. This will be the drain whenthe mold is ready. The ring must be strong enough tohold the rock, which is placed on top of it, in its place(see fig.’1). Now we grease the inside of the box and therock with soft soap. This will allow us to release the castwhen it has hardened.

To be able to dismantle the mold in order to remove

the rock (and later the cast rock), it will be made of sev-eral layers. Every layer must harden before a new one isapplied. The number and thickness of the layers is de-pendent on the shape and the size of the rock (see fig. 2).Prepare enough plaster to make the first layer. The plas-ter must have the consistency of yoghurt and be clearedof any dry clumps of gypsum. When the plaster is hardenough, cut some grooves in the surface. These will be

Fig 1. The rock is placed on a 3 cm-high ring of clay.

Fig 2. This mold consists of five parts.

Fig 3. The mold is filled to the brim with the casting clay.

Make your own reefGerard Tijsseling

filled with the next layer and help fixing the differentlayers when they are hardened. Before we pour the nextlayer the surface of the previous is greased with soft soap.A ring of clay is placed on top of the rock before the lastlayer is poured. This ring has a diameter of about 8 cmand will later be the opening through which the mold isfilled.

When the plaster of the last layer is strong enough totake it apart, all layers are removed and the soaped washedof. Sometimes, depending on the shape of the rock, alayer has to be sawed in two (see fig. 2). Now we dry allparts of the mold for several weeks.

Casting the rock

For making the rocks we need casting-clay which canbe bought or made by mixing water with clay powder. Itmust have the consistency of yogurt.

The mold is reconstructed – if we have made somegrooves it will hold the layers perfectly in place – andfixed by rubber bands. In this case, the inner tube of abicycle wheel works fine. We close the drain with a

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Fig 4. Care must be taken when dismantling the mold.

Fig 5. The cast is trimmed to obtain a smooth surface.

Fig 6. One or two larger holes are made to enable entrance.

Fig 7. Small holes in the cast release trapped air.

Fig 8. A completely ceramic reef in the author’s

All photos by Iemkje Tijsseling

wooden stopper or of some other material which doesnot absorb water. The mold is now completely filled withthe fluid clay (see fig. 3).

The water in the casting-clay is absorbed by the plas-ter of the mold. Every five to ten minutes we top of theclay. After a half, with very dry molds, to one hour wedrain the remaining clay in a bucket (it can be re-used).Now a layer with a thickness of about one centimeterlines the mold inside. We let this dry for about two tothree days before we dismantle the mold (see fig. 4). Thefilling opening and the drain in the cast are closed with

soft, moist clay. Any irregularities, caused by the struc-ture of the mold, are trimmed from the cast (see fig. 5).

When we have decided what side will be facing theback of the aquarium, we cut here one or two holes topermit entrance by the fish (see fig. 6).

Also some smaller holes are punctured on all sides ofthe rock in order to release the air that got trapped inside(see fig. 7). When you forget these small holes, you willfind your carefully constructed reef floating by when theaquarium is filled. With some artistic experience a castcan be re-shaped a little by denting it. This is, however,not mandatory. It is remarkable how many identical

“rocks” one can place in an aquarium without giving theimpression that all “rocks” are the same.

After the casts have dried for several weeks they haveto be baked in a kiln at 1000 to 1200°C, depending onthe type of clay. A friendly potter or brick factory mighthelp you out if you have no ready access to such an oven.The color of the clay or glazing is of minor importancebecause the ceramic rocks will be covered with algae ina few month’s time (see fig. 8).

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The chief problem in regularly breeding Julidochromisspecies or indeed any of the small rock dwellingTanganyikans results from the difficulty in removingthe young fish without disturbing the parents. The usualmethod involves removing all rockwork from theaquarium before it is possible to net the young fishout. The rocks are then replaced causing great upheavaland stress to the parents which usually shows itself bytheir unwillingness to spawn again until they have set-tled down, which can take as much as 6 to 8 weeks.Also, when subjected to this amount of disturbance, itsometimes happens that the pair “fall out” with eachother to the extent that one may be actually killed.

Re-pairing adult Julidochromis and some other cave-inhabiting cichlids is something that I now never do,as the pair bond is never as strong as with their origi-nal mates, so there is a tendency for intermittent fight-ing. It is far better to start again with a selection ofyoung fish to give a choice of mates and wait for astable pair to form.

My method for maintaining and breeding these fishremoves most of these problems.

I have found it best to custom build a doubleaquarium to accommodate two pairs of fish. First, builda basic aquarium 48 x 18 x 18 inches (122 x 45 x 45cm) – 6 mm glass is adequate for this as the shelvesand divider strengthen it and stop it flexing when full.A clear divider glass is siliconed in place to divide theaquarium into two equal halves, approximately 24 x18 x 18 inches (60 x 45 x 45 cm). The seal on thismust be watertight.

A glass shelf measuring 24 x 6high x 9wide (60 x13 x 22’cm) is then fitted along the back of each com-partment. This should be made separately (6’mm glass)and painted on what will be the underside with two orthree coats of matt black or any dark coloured paint.The shelf should appear “solid” when finished. Thedark colour enhances the markings on Julidochromisand helps to prevent them looking “washed out” aswell as discouraging algae growth. The corners of theshelf should be rounded off with a grinding disc inorder to accommodate the inner silicon seal of theaquarium.

It is absolutely vital to make certain that all seals onthe shelf are watertight, forming an airtight compart-ment under the shelf. If water should seep into thiscompartment it will make the paint fall off the under-side of the shelf. Removing it for repainting is verydifficult indeed (I speak from experience here!) and itis far easier to make certain you have a good seal in

A Julidochromis breeding tank

John Szwechlowicz

the first place.When the aquarium is ready for use, rocks and/or

clay flowerpots are arranged to form the usual caveson top of the shelf. No substrate is necessary and fil-tration is provided by a double Algarde Biofoam 200or similar foam filter. It is a good idea to direct thefilter outlet towards the back of the tank to help pre-vent sediment settling among the rocks. It will beflushed out to collect in the front of the aquarium fromwhere it can be periodically syphoned out. Mainte-nance is thus greatly simplified.

A pair of Julidochromis (or other rock dwellingcichlids) is placed in each side of the aquarium. Herethey will live and breed as in a conventional aquarium,but several benefits soon become apparent. Becauseeach pair can see another pair on the very edge of theirterritory, much time is spent threatening the oppositepair, although the divider prevents actual physical con-tact. This tends to occupy their minds and dramati-cally reduces the chances of a pair falling out witheach other, which I consider to be mainly due to thefishy equivalent of boredom, when they are not hav-ing to actively defend a territory.

If good water quality is maintained, sooner or laterthe pair will produce fry. Initially, these stay in amongthe rocks as normal but as they reach about 2 or 3 cmin length, the parents evict them from the rocks, downinto the front of the aquarium where they tend to con-gregate around the sponge filter as it is the only shel-ter in that part of the tank. At this stage they should beremoved as the parents will be preparing to spawnagain. This is accomplished by syphoning the waterout into a bucket, down to the level of the top of theshelf. Some of this water is then poured over the backof the rocks with a jug in order to flush out any frywhich darted back into the rocks due to the initial dis-turbance. The fry are then easily removed as there isnowhere to hide and not enough water to enable themto get back into the rocks. It does not matter too muchif the adults are flushed out at the same time, provid-ing they are not netted out and the positions of therocks remain absolutely untouched, i.e. their territoryis unchanged. The water is then returned to the tank.

Since the adults territory is exactly the same, therocks not having been moved at all, and they also havethe opposite pair to distract them, they tend to settledown again very quickly, with a much reduced chanceof any fighting. If a 25 to 30% waterchange is doneabout two days later, it is usually enough to triggeranother spawning and the sequence is repeated

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Basic aquarium 48x18x18 inches(122x45x45 cm) shown with clearglass divider already siliconed inplace

painted onunderside

all corners ground offto avoid any damage toinner seal of aquarium

shelves are assembledready to be sealed into

the aquarium

place rockworkon top of shelf

airtightcompartment

Above: The construction of a specialist aquarium for breeding Julidochromis and similar species.Below: The aquarium with the shelves installed. Drawings by John Szwechlowicz.

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The family Cichlidae is exceptionally rich in species. Ofcourse, this fact is caused by the ease with which thesefishes are able to speciate. Moreover, cichlids are alsoknown for their plasticity, which means that within onespecies considerable phenotypic variation can occur. Itis more than justified when the South African ichthyolo-gist Michael Bruton (1989) states: “The adoption of al-ternative phenotypic states in nature is probably morewidespread than is currently realised and manypopulations which are currently recognised as speciesmay in fact, be no more than ecophenotypes of one oranother homeorhetic state”. It might thus be clear thatthe question whether one or another population or groupof populations should be regarded as a distinct species ornot, will be a hotly debated issue among cichlid scien-tists for many years to come.

Evolutionary biologists usually call attention to thecichlids of Lake Nabugabo whenever they want to dem-onstrate that speciation in these fishes can take place in arelatively short period of time. Lake Nabugabo was partof Lake Victoria in previous times. About 4000 yearsago a bay was separated from the main lake by a sandbar.In the lake thus formed Greenwood (1965) recognizesfive distinct species which must have developed fromancestral Victorian populations which remained in theisolated bay. Since Greenwood’s paper the cichlids ofLake Nabugabo have been presented in many publica-tions as examples of how much faster speciation mightprogress than was previously accepted.

Some recent observations, however, suggest thatspeciation among cichlids may operate at an even fasterspeed than the cichlids of Lake Nabugabo let us assume.One of these observation relates to the description of anew Mexican cichlid (Werner & Stawikowski, 1988).The new species, which is named Paratheraps breidohriby the authors, is reported to be endemic to Presa de laAngostura, a reservoir in the southern part of Mexico.The closest relative of P. breidohri, named Paratherapshartwegi by the same authors, occurs in the rivers andstreams flowing into the reservoir as well as in the reser-voir itself. Hence, we cannot escape the notion that P.breidohri developed from a population of P. hartwegiafter the man-made dam formed the reservoir. This wouldmean that P. breidohri, if accepted as a distinct species,would just be less than 30 years old! On the other handwe cannot exclude the possibility that P. breidohri shouldbe regarded as the lacustrine morph of P. hartwegi. Fur-ther examination is required to provide a solution to thesequestions.

Very recently it became apparent that human inter-vention may have yielded a new species in Africa as well.This species was named Haplochromis erythromaculatus

by the authors, De Vos et al. (1990). H. erythromaculatusis found in Lake Bulera and Lake Ruhondo. These lakeslie in the northern part of Rwanda where, as early as1907, the German H. Schubotz investigated the fish faunaof Lake Ruhondo. He concluded that this lake was in-habited only by small barbs, but certainly did not men-tion cichlids. Later fishery biologists tried to enrich thefish fauna of the Rwandan lakes with species suitable forconsumption. Therefore they introduced several tilapiineswhich were collected in Lake Edward. Among the tilapiasthere were apparently also some specimens of ahaplochromine. From these “contaminating” individu-als H. erythromaculatus must have been originated (in aperiod of time which is less than 80 years). De Vos et al.tried to discover what species in Lake Edward was trans-ferred to Lake Bulera and Lake Ruhondo, but could notfind it. The cichlids of Lake Edward are apparently notwell studied yet. Nevertheless, the recognition of H.erythromaculatus as a valid species suggests that cichlidsare able to speciate much faster than previously thoughtby evolution biologists. Jos Snoeks (pers. comm.), oneof the authors, believes that speciation needs more timethan the suggested 80 years.

Not only do human interventions in nature allow us tostudy the speed of speciation among cichlids, but alsoour expanding knowledge about the ecological historyof the waters in which cichlids live contributes to thesestudies. Moreover such knowledge suggests thatspeciation among cichlid can take place at a much higherspeed than is generally accepted.

Recently, Owen et al. (1990) showed that the level inLake Malawi fluctuated more frequently and betweenmuch higher extremes (not the yearly fluctuations) thanhad been concluded from previous studies. From thepaper by Owen et al. it appears that the level of LakeMalawi, in the last decades of the eighteenth century andin the first decades of the nineteenth century, was at least120 meters below that of today’s lake. If these findingsprove to be true it would mean that about 150 to 250years ago many of the locations which are presently in-habited by endemic Mbuna were a part of the continent.This is true for the islands Likoma, Chizumulu, Mbenji,the Maleris, Boadzulu, Thumbi West, etcetera. The au-thors conclude that the Mbuna, which are endemic tothese islands, cannot be older than the mentioned 150 to250 years (see editor’s note). For the time being we re-gard these endemic species as valid species and not asecophenotypes.

Now, examining the bathymetric map of Lake Ma-lawi, we should be able to indicate from which locationsthe endemic species of certain islands were derived andwhich species can be regarded as their ancestors.

The last minutes of speciation

Martin Geerts

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In the frame of this yearbook it is, unfortunately, notfeasible to continue the discussion of these examples orto detail others. Cichlidists with a multidisciplinary back-ground will, without doubt, be able to find more andmaybe better examples. Then it will be possible to gain abetter understanding of the process which is calledspeciation, of which it is usually thought that a man’slifetime is too short to see it happen.

References

BRUTON, M. (1989) The ecological significance of alternativelife-history styles. pp. 503-553 in M. Bruton (ed.) Alterna-tive life-history styles of animals. Dordrecht, Netherlands.

DEVOS, L., SNOEKS, J. & D.F.E. THYS V. D. AUDENAERDE (1990)Description d’Haplochromis erythromaculatus, especenouvelle (Teleostei, Cichlidae) des Lacs Bulera et Ruhondo,Rwanda. Ichthyol. Explor. Freshwaters Vol. 1 (3); pp 257-268.

GREENWOOD, P.H. (1965) The cichlid fishes of Lake Nabugabo,Uganda. Bull. Br. Mus. nat. hist. (zool), 12; pp 315-357.

OWEN, R.B., R. CROSSLEY, T.C. JOHNSON, D. TWEDDLE, I.KORNFIELD, S. DAVISON, D.H. ECCLES & D.E. ENGSTROM,(1990) Major low levels of Lake Malawi and their impli-cations for speciation rates in cichlid fishes. Proc. R. Soc.Lond. B-240; pp 519-553.

WERNER, U & R. STAWIKOWSKI (1988) Ein neuer Buntbarschaus Südmexico: Paratheraps breidohri gen. nov. spec. nov.DATZ 41 (1); pp 20-23.

Editor’s note

In the paper of Owen et al. (1990) it is concluded that mostof the endemic species inhabiting islands, which were partof the mainland 200 years ago, have developed in recenttimes. Disregarding the fact that the definition of a speciesis still hotly debated and will probably never become auniversally agreed entity, there are some species (distincttaxa) which are found on both sides of the once dried-upsouthern part of Lake Malawi. This would suggest that, duringthe low lake level, there must have been a suitable habitatwhere at least these species could have survived the“drought”. These species are P. barlowi (distribution: Mbenji,Maleris, Thumbi West, Nkhudzi and Eccles Reef), P. sp.“Dumpy” (distribution: Maleris and Fort Maguire), P. sp.“Zebra Red Dorsal” (distribution: Mpanga Rocks,Nakantenga, Nkudzi and Eccles Reef) and Copadichromisazureus (distribution: Nkhomo Reef, Mbenji, Maleris andEccles Reef).One of the habitats that could have served as a “sanctuary”during a low lake level is a reef between Chinyamwezi Islandand the Nankhumba peninsula. This reef remains at least24 meters below the surface while its basis meets the sandat a depth of at least 105 m. I have visited this reef inDecember 1990 and found Cynotilapia sp. “Chinyankwazi”in vaste numbers (see photo). Also present were P.tropheops, which is further distributed around theNankhumba peninsula and Chinyamwezi, P. flavus, whichis known from Chinyankwazi and P. ater, known fromChinyankwazi and Chinyamwezi.

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GOLDSCHMIDT, T. & J. DE VISSER (1990) On the pos-sible role of egg mimics in speciation. ActaBiotheoretica 38: pp 125-134.

The potential mechanisms of speciation in themouthbrooding haplochromine cichlids is a topicthat has been widely studied and hotly debated inrecent times. There have been numerous pro andcon arguments presented in regard to the possiblespeciation effects of both the courtship behavior andmating system of these fishes. One interesting ap-proach to this topic is presented in this paper. While,like most such studies, it is highly theoretical in itsapproach, the authors feel that sufficient evidenceis at hand to warrant both a preliminary report andfurther pursuit of the topic.

This study is based on initial observations of aLake Victorian species Haplochromis “argens”.This species is widespread in Mwanza Gulf, but inEmin Pasha Gulf what is described as a “....verysimilar “species” or variety of H. “argens”....” wasfound mixed in with catches of H. “argens”. Thisfish was designated as H. dusky “argens”. In pre-liminary studies, females of the two fishes couldnot be distinguished, but the males were noticeablydifferent. The males differed not only in body color(H. “argens” = silver white with red caudal; H.dusky “argens” = dusky grey with no red on cau-dal) but also in the number, color and placement ofthe egg dummies on the anal fin. Based on this ini-tial observation and other information at hand, theauthors present two theories (models) by which eggdummies and their potential subsequent divergencein a species may trigger reproductive isolationwhich in turn could lead to the formation of newspecies.

The first model (with the proposed designationof mimetic isolation) supposes that if there arechanges in the characteristics of the eggs (size,shape) in small temporarily isolated populations thatmatching changes in the egg dummies will also de-velop in the males. This model stipulates that thiswill be for species in which the egg dummies accu-rately mimic the eggs. Should these factors takeplace, the fish involved would then be repro-ductively isolated, if re-united with its former largerpopulation. This is noted to be only a theoreticalmodel and that no direct positive evidence is avail-able to substantiate its validity.

The second model, again dealing with small tem-porarily isolated populations, involves initial

Eggspots and Ocelli

Reviews by Lee Finley

changes in the egg dummies based on selection pres-sures aimed at them. Such pressures may includehabitat (light vs. dark areas) and potential for pre-dation. The authors note some evidence generallyindicating that cichlids living in darker areas tendto have more numerous, larger, or more colorful(“supra-normal”) egg dummies when compared tothose living in lighter areas. Such adornment in lightareas might have poor selection factors by makingthe fish more visible to predators. Again, if suchchanges did take place these fish would bereproductively isolated should they rejoin theiroriginal population. Although this model is, like thefirst, lacking in proper testing, some field evidencedoes tend to support it. The authors also suggestthat laboratory studies may be worthwhile in shed-ding some light in this area.

In this paper the authors also provide a handylisting of the possible functions of egg dummies andnote that the function (or functions) of them in spe-cies to be studied “......should preferably be known”.

WINEMILLER, K. D. (1990) Caudal eyespots as de-terrents against fin predation in the neotropicalcichlid Astronotus ocellatus. Copeia (3): pp 665-673.

The variety of colors and patterns observed incichlids are fascinating and may often lead one intoquestioned thought as to why they are the way theyare. This paper attempts to answer that question forone species of cichlid, the so-called Oscar.

Most aquarists are familiar with the Oscar andits distinctive color pattern. Most noticeable is thelarge ocelli, or “eyespot” that is located near thebase of the caudal fin. It is this feature and its pos-sible function that is addressed in this paper. Theo-ries regarding the function of ocelli in cichlids arecertainly not new. In 1977 T. M. Zaret presented astudy on another cichlid, Cichla ocellaris. Zaret’sconclusion was that the ocelli serve as a form ofspecies recognition which inhibit cannibalism. Fur-thermore, according to Zaret, A. ocellatus (and othercichlids such as Crenicichla spp.) have evolvedmimic ocelli to avoid predation by the Cichla.

Winemiller possesses a different vision in regardto the possible function of the ocelli in the Oscar.Based on the results of a field study in Venezuelathat spreads over five years, the author has con-

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Some West African cichlids show other types of spots in their fins. Do they have a function as well? Photo by R. Numrich.

cluded that the ocelli acts as an eye mimic with thefunction being to reduce the attacks of fin eatingpiranha (mainly of the genus Serrasalmus).

A. ocellatus, and the co-existing cichlid Caque-taia kraussii (which also possesses ocelli), werecollected throughout the year and comparisons weremade as to the extent of fin predation by piranha.For the four driest months of the year, when pira-nha were absent from the study area, no damagewas noted. But for the rest of the year the resultswere quite dramatic. When compared in a series ofsize groupings, the C. kraussii always showed moreextensive damage than that seen in A. ocellatus. Theauthor found this interesting in that considering thefeeding and behavioral styles one would expect re-verse results. C. kraussii is decribed as a “sit andwait” predator and theoretically would have moreopportunity to “...detect and avoid.....” piranha. Onthe other hand A. ocellatus spends much of its timeslowly swimming and examining the roots of float-ing plants searching for its prey which consists ofalmost 60% insects (both aquatic and terrestial).

From the above, the author concludes that theocelli of A. ocellatus, which resembles the eye bothin size and color (as compared to that of C. kraussii

which does not), combines with the anal, dorsal andcaudal fin shapes to form a full caudal mimic of thehead region which thereby reduces or prevents at-tacks by fin eating piranha. Additional backup theo-ries are also presented (the fins don’t taste good;Oscars as social fishes may be better able to fendoff attacks than the solitary C. kraussii, etc.).

Supporting aquarium studies using fin eating pi-ranha and a variety of fishes, including ocellatedand non-ocellated cichlids, show that ocellatedcichlids fare the best by far. One last, and very in-teresting experiment was attempted. Both A.ocellaris and C. kraussii were set up in concretepools with piranha. Grease paint was used to ob-scure and accentuate the ocelli of both species.Unfortunately, before observations could be madeas to the effects of this, the paint “wore off”. It isnoted that similar experiments would prove usefulfor further testing the “...fin-predator/head mimicryhypothesis”.

98

AustraliaThe New South Wales Cichlid Society

P.O. Box 163Moorebank, N.S.W. 2170

Queensland Cichlid GroupP.O. Box 163

Wooloongabba, Queensland 4102

Victorian Cichlid Society23 Mangana Drive

Mulgrave, Victoria 3170

AustriaDeutsche Cichliden GesellschaftVictor Kaplan Straße 1-9/1/3/12

A-1220 Wien

BelgiumBelgische Cichliden Vereninging

Kievitlaan 23B-2228 Ranst

CzechoslovakiaSZCH Klub Chovatelov Cichlíd

Príkopova 2CS-831 03 Bratislava

DenmarkDansk Cichlide Selskab

Ornevej 58, st. tv.DK-2400 Kobenhavn NV

FranceAssociation France Cichlid

15 Rue des HirondellesF-67350 Dauendorf

GermanyDeutsche Cichliden Gesellschaft

Eberescheweg 41D(W)-4200 Oberhausen

NetherlandsNederlandse Cichliden Vereniging

Boeier 31NL-1625 CJ Hoorn

SwedenNordiska CiklidSällskapet

Skogsgläntan 16S-435 38 Mölnlycke

CICHLID ORGANIZATIONS WORLDWIDE

SwitzerlandDeutsche Cichliden Gesellschaft

Am Balsberg 36CH-8302 Kloten

United KingdomBritish Cichlid Association

100 Keighley RoadSkipton, North Yorkshire, BD23 2RA

U. S. A.American Cichlid Association

P.O. Box 32130Raleigh, NC 27622

Apistogramma Study Group1845 Jaynes Road

Mosinee, WI 64455

Cichlasoma Study Group6432 South Holland Court

Littlerton, CO 80123

Atlantic Cichlid & Catfish Organization29 Pearsall Avenue

Jersey City, NJ 07305

The Greater Chicago Cichlid Association5712 South Drexel 1Chicago, IL 60637

Illinois Cichlids and Scavengers7807 Sunset Drive

Elmwood Park, IL 60635

Michigan Cichlid AssociationP.O. Box 59

New Baltimore, MI 48047

Pacific Coast Cichlid AssociationP.O. Box 28145

San Jose, CA 95128

Southern California Cichlid Association1610 East McFaddenSanta Ana, CA 92705

Texas Cichlid Association6845 WinchesterDallas, TX 75231

99

Introduction ................................................................ ................................................................................... 3

Tanganyikan CichlidsThe giant featherfin by René Krüter .......................... ................................................................................... 4Three beautiful sanddwelling cichlids ....................... ................................................................................... 7Neolamprologus sp. “Cygnus” .............................. 11 Petrochromis, three new variants ........................... 20Altolamprologus calvus ......................................... 12 Tropheus moorii ...................................................... 21Neolamprologus sexfasciatus ................................ 13 Cyathopharynx furcifer ........................................... 22Neolamprologus leloupi ......................................... 14 Xenotilapia flavipinnis ............................................ 23Neolamprologus mustax......................................... 15 Xenotilapia papilio ................................................. 24Greenwoodochromis christyi ................................. 16 Xenotilapia sp. “Katete” ......................................... 25Gnathochromis permaxillaris ................................ 17 Cyprichromis sp. “Leptosoma Jumbo” .................. 26Trematocara nigrifrons .......................................... 18 Paracyprichromis nigripinnis ................................ 28Microdontochromis tenuidentatus ......................... 19

Malawian CichlidsThe blue sanddwellers ............................................... ................................................................................. 29Aulonocara auditor (Trewavas, 1935) ...................... ................................................................................. 33The genus Tyrannochromis Eccles & Trewavas by Ethelwynn Trewavas ........................................................ 36Maravichromis (Caprichromis) liemi .................... 40 Copadichromis sp. “Virginalis Chitande” .............. 47Sciaenochromis gracilis ......................................... 41 Diplotaxodon sp. “Big Head”................................. 48Otopharynx decorus ............................................... 42 Lethrinops sp. “Christyi Fort Maguire” ................. 49Maravichromis sp. “Double Spot” ........................ 43 Taeniolethrinops sp.”Furcicauda Ntekete” ............ 50Nyassachromis sp. “Mpanga” ................................ 44 Lethrinops sp. “Longipinnis Ntekete” ................... 51Otopharynx (Ctenopharynx) nitidus ...................... 45 Gephyrochromis sp. ................................................ 52Copadichromis sp. “Virginalis Gome” .................. 46 Pseudotropheus sp. “Zebra Charo” ........................ 53

Victorian CichlidsExciting new discoveries by Laif DeMason .............. ................................................................................. 54

West African CichlidsTwo new Chromidotilapia-species by Roland Numrich 57Steatocranus ubanguiensis/Steatocranus mpozoensis61Pelvicachromis taeniatus ........................................... ................................................................................. 62

Central American CichlidsThe re-discovery of Paraneetroplus nebuliferum by Juan Miguel Artigas Azas ............................................. 64“Cichlasoma” spinosissimum (Vaillant & Pellegrin, 1902) by Willem Heijns ................................................ 68“Cichlasoma” melanurum..................................... 71 “Cichlasoma” bartoni ............................................ 74“Cichlasoma” sp. “Pantepec” ............................... 72 “Cichlasoma” labridens ........................................ 75“Cichlasoma” sp. “Poza Rica” ............................. 73

South American CichlidsTahuantinsuyoa macantzatza Kullander, 1986 by Ron Bernard ....................................................................... 76Biotodoma sp. “Santarém” by Ron Bernard .............. ................................................................................. 78Retroculus lapidifer (De Castelnau, 1855) by Ron Bernard ............................................................................. 80Some large Crenicichla by Frank Warzel .............. 82Crenicichla compressiceps .................................... 86 Symphysodon aequifasciatus .................................. 88Crenicichla cyclostoma .......................................... 87

Cichlid MaintenanceMake your own reef by Gerard Tijsseling................. ................................................................................. 90A Julidochromis breeding tank by John Szwechlowicz ................................................................................. 92

Cichlid LiteratureThe last minutes of speciation by Martin Geerts ...... ................................................................................. 94Eggspots and ocelli by Lee Finley ............................. ................................................................................. 96

The CICHLIDS yearbookCONTENTS

100 ISBN 3-928457-00-4

Gnathochromis permaxillaris, one of the most interesting cichlids from Lake Tanganyika.

A female Crenicichla marmorata from Santarém. Photo by Frank Warzel.

The CICHLIDS yearbook